Remotely sensed variability of temperature and chlorophyll in the southern Benguela: upwelling frequency and phytoplankton response

High-resolution (1km) satellite data from the NOAA AVHRR (Advanced Very High Resolution Radiometer) and OrbView-2 SeaWiFS (Sea-viewing Wide Field-of-view Sensor) are used to investigate the upper layer dynamics of the southern Benguela ecosystem in more detailed space and time scales than previously undertaken. A consistent time-series of daily sea surface temperature (SST) and chlorophyll a concentration images is generated for the period July 1998–June 2003, and a quantitative analysis undertaken. The variability in SST, upwelling and phytoplankton biomass is explored for selected biogeographic regions, with particular focus on intra-seasonal time scales. The location and emergence of upwelling cells are clearly identified along the length of the southern Benguela, being distinct on the narrow inner and the mid-continental shelves. Most notable is the rapidly pulsating nature of the upwelling, with intense warm/cold events clearly distinguished. The phytoplankton response to this physical forcing is described. Chlorophyll concentration on the inner shelf largely mirrors the pattern of SST variability, similarly dominated by event-scale processes. Over the mid-shelf, higher chlorophyll is observed throughout all seasons, although low biomass occurs during winter. The variability of the offshore extent of SST and chlorophyll is identified at locations of differing shelf width. Cooler upwelled water is confined primarily to the narrow inner-shelf, with event-scale pulses extending considerable distances offshore. Agulhas Current influences are readily observed, even on the Cape Peninsula inner-shelf. Chlorophyll concentrations vary considerably between the locations of differing shelf width. SST, upwelling and phytoplankton indices are derived for selected locations to quantify the intra-seasonal variations. The SST indices show marked temperature changes associated with rapid pulsation on the event scale. No strong seasonal signal is evident. In contrast, the upwelling indices display a strong seasonal signal, with most intense upwelling occurring in spring/summer in the south. The phytoplankton response to the seasonal upwelling index differs between the selected locations. This study concludes that, although low-resolution SST and chlorophyll data may be useful for investigating general patterns over large scales, higher resolution data are necessary to identify finer scale spatial and temporal variability, especially in the inshore coastal zones.     

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.     

Influence of buoyancy and vertical distribution of sardine Sardinops sagax eggs and larvae on their transport in the northern Benguela ecosystem

In recent years, sardine Sardinops sagax spawning has been recorded inshore off central Namibia. Field observations on eggs and laboratory measurements show that spawning, demonstrated by the distribution of newly spawned eggs, takes place just below the upper mixed layer. The high positive buoyancy of the eggs causes them to ascend rapidly to the surface layer, where they are moved offshore by upwelling-induced offshore transport. However, increased wind-induced mixing also influences the vertical distribution of eggs, causing them to be partly mixed down below the layer moving offshore and into the layer moving inshore. This mechanism acts to retard the transport and offshore loss of eggs from the spawning areas. The vertical distribution of sardine larvae, with highest concentrations deeper than 20 m, indicates active movement out of the layer moving offshore, and this tendency seems to be more pronounced for older larvae. Hence, vertical migration of larvae is an additional factor mitigating their loss from nearshore. Taken together, these features seem to minimize the offshore loss of offspring, particularly in periods of low stock biomass when spawning close to the shore seems to be common.     

Wintertime rates of net primary production and nitrate and ammonium uptake in the southern Benguela upwelling system

The elevated levels of primary productivity associated with eastern boundary currents are driven by nutrient- rich waters upwelled from depth, such that these regions are typically characterised by high rates of nitrate-fuelled phytoplankton growth. Production studies from the southern Benguela upwelling system (SBUS) tend to be biased towards the summer upwelling season, yet winter data are required to compute annual budgets and understand seasonal variability. Net primary production (NPP) and nitrate and ammonium uptake were measured concurrently at six stations in the SBUS in early winter. While euphotic zone NPP was highest at the stations nearest to the coast and declined with distance from the shore, a greater proportion was potentially exportable from open-ocean surface waters, as indicated by the higher specific nitrate uptake rates and f-ratios (ratio of nitrate uptake to total nitrogen consumption) at the stations located off the continental shelf. Near the coast, phytoplankton growth was predominantly supported by ammonium despite the high ambient nitrate concentrations. Along with ammonium concentrations as high as 3.6 µmol l^–1, this strongly suggests that nitrate uptake in the inshore SBUS, and by extension carbon drawdown, is inhibited by ammonium, at least in winter, although this has also been hypothesised for the summer.     

Thermal constraints on the respiration and excretion rates of krill,Euphausia hanseni and Nematoscelis megalops,in the northern Benguela upwelling system off Namibia

Rates of respiration and ammonia excretion of Euphausia hanseni and Nematoscelis megalops were determined experimentally at four temperatures representative of conditions encountered by these euphausiid species in the northern Benguela upwelling environment. The respiration rate increased from 7.7 µmol O₂ h^?1 g_ww ^?1 at 5 °C to 18.1 µmol O₂ h^?1 g_ww ^?1 at 20 °C in E. hanseni and from 7.0 µmol O₂ h^?1 g_ww ^?1 (5 °C) to 23.4 µmol O₂ h^?1 g_ww ^?1 (20 °C) in N. megalops. The impact of temperature on oxygen uptake of the two species differed significantly. Nematoscelis megalops showed thermal adaptations to temperatures between 5 °C and 10 °C (Q₁₀ = 1.9) and metabolic constraint was evident at higher temperatures (Q₁₀ = 2.6). In contrast, E. hanseni showed adaptations to temperatures of 10–20 °C (Q₁₀ = 1.5) and experienced metabolic depression below 10 °C (Q₁₀ = 2.6). Proteins were predominantly metabolised by E. hanseni in contrast to lipids by N. megalops. Carbon demand of N. megalops between 5 and 15 °C was lower than in E. hanseni versus equal food requirements at 20 °C. It is concluded that the two species display different physiological adaptations, based on their respective temperature adaptations, which are mirrored in their differential vertical positioning in the water column.     

On vertical velocity fluctuations and internal tides in an upwelling region off the west coast of India

Hourly fluctuations of vertical velocity in relation to components of flow and wind and temperature oscillations at a morring site in the shelf waters off the west coast of India are discussed. The vertical velocities were computed from a time series of vertical temperature profiles assuming that horizontal advection of temperature is negligible. The computed values at a depth of 40 m during the 72-h period of observation were of the order of 10⁻¹ to 10⁻²cm s⁻¹, with a mean value of −2·77 × 10⁻² cm s⁻¹ indicating a net upward movement of water. The computed vertical velocity showed fluctuations of about 2–3 h, in addition to weaker signals of about 12 h. Based on the spectral estimates, we speculate that these fluctuations of 2–3 h in the vertical velocity may be caused by the fluctuations in the along-shore wind. The oscillations of isotherms found in the temperaturedepth time series and the spectral estimates of temperature and cross-shore flow component showed a periodicity of about 12 h, which indicated the presence of semi-diurnal internal waves. The fact that these internal wave troughs were associated with the measured onshore flow suggested that the waves were propagating offshore. The computed stability parameters showed little evidence of instability or mixing. It was found that the isotherm troughs in the temperaturedepth time series at about 12-h period coincided with high vertical shear in the cross-shore direction and low values of Brunt Vaisälä frequency.     

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.     

Partitioning of a goby resource amongst four avian predators and evidence for altered trophic flow in the pelagic community of an intense,perennial upwelling system

Pelagic gobies Sufflogobius bibarbatus were numerically the most important prey of jackass penguins Spheniscus demersus. Cape cormorants Phalacrocorax capensis and bank cormorants P. neglectus sampled at islands off the South West African coast during the period 1978–1982. These three seabirds feed on small gobies near the surface and some of them can also dive sufficiently deep to catch larger gobies. Their populations at islands north of Lüderitz, where gobies are abundant, have been increasing. Cape gannets Morus capensis feed only on large gobies that are infrequently at the surface, and the gannet population off South West Africa has shown a large decrease since the collapse of the pilchard Sardinops ocellata resource in the late 1960s and early 1970s. In the intense perennial upwelling system situated between 22 and 27°S gobies are believed to have partially replaced pilchards during the 1970s. Both pilchards and gobies are able to feed on large diatoms of the genera Chaetoceros and Delphineis, which dominated the inshore phytoplankton in the early 1970s when biomass levels of both pilchards and gobies were low.