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.     

Abrupt environmental shift associated with changes in the distribution of Cape anchovy Engraulis encrasicolus spawners in the southern Benguela

Cape anchovy Engraulis encrasicolus spawners in the southern Benguela showed an eastward shift in their distribution on the Agulhas Bank that occurred abruptly in 1996 and has since persisted. We assessed whether this shift was environmentally mediated by examining sea surface temperature data from different regions of the Agulhas Bank, which showed that in 1996 the inner shelf of the Agulhas Bank to the east of Cape Agulhas abruptly became 0.5°C colder than in previous years and has since remained that way. In addition, signals, coherent with the 1996 shift recorded in sea surface temperatures, were also found in atmospheric surface pressure and zonal wind data for that region; interannual coastal SST variability is also shown to be correlated with zonal wind-stress forcing. As a result, increased wind-induced coastal upwelling east of Cape Agulhas is proposed as the main driver of the observed cooling in the coastal region. The synchrony between the environmental and biological signals suggests that the eastward shift in anchovy spawner distribution was environmentally mediated and arose from a change in environmental forcing that altered the relative favourability for spawning between regions to the west and east of Cape Agulhas. The results highlight how a relatively minor change in environmental conditions can lead to a drastic spatial reorganisation of the life history of one species in an ecosystem.     

Characterising and comparing the spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem were characterised by comparing their egg abundances with environmental variables measured concomitantly during two different survey programmes: the South African Sardine and Anchovy Recruitment Programme (SARP), which comprised monthly surveys conducted during the austral summers of 1993/94 and 1994/95; and annual pelagic spawner biomass surveys conducted in early summer (November/December) from 1984 to 1999. Eggs were collected using a CalVET net. Physical variables measured included sea surface temperature (SST), surface salinity, water depth, mixed-layer depth, and current and wind speeds; biological variables measured included phytoplankton biomass, and zooplankton biomass and production. Spawning habitat was identified by construction of quotient curves derived from egg abundance data and individual environmental variables, and relationships between these variables were determined using multivariate co-inertia analysis. SARP data showed that anchovy spawning was associated with cool water and moderate wind and current speeds, whereas sardine spawning was related to warmer water and more turbulent and unstable conditions (i.e. high wind speeds and strong currents) than for anchovy. SARP data also showed significant differences in selection of spawning habitat of the two species for all environmental variables. The relationship between anchovy egg abundance and salinity was strongly positive, but strongly negative with water depth, phytoplankton biomass and zooplankton production. Sardine egg abundance was strongly positively related to current speed. The spawner biomass survey data demonstrated that the spawning habitat of anchovy was characterised by warm water and high salinity, whereas sardine spawning was associated with cool water and low salinity. The survey data showed significant differences in spawning habitat selection by anchovy and sardine for SST, salinity and zooplankton biomass, but not for the other environmental variables. There was a positive relationship between anchovy egg abundance and SST, salinity and mixed-layer depth, and a negative relationship with water depth, phytoplankton biomass and zooplankton production. For sardine there was a strong positive relationship between egg abundance and current speed and wind speed. Differences in the results between the two survey programmes could be attributable to differences in their spatio-temporal coverage. Spawning habitats of anchovy and sardine appear to be substantially different, with anchovy being more specific than sardine in their preference of various environmental conditions.     

Diet of sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The diet of sardine Sardinops sagax in the southern Benguela was investigated by microscopic examination of stomach contents. The relative dietary importance of prey size and prey type was assessed by calculating the carbon content of prey items. Sardine is an omnivorous clupeoid, ingesting both phytoplankton and zooplankton, with the relative importance of these two food types varying both spatially and temporally. Stomach contents were numerically dominated by small prey items, principally dinoflagellates, followed by crustacean eggs, cyclopoid copepods, calanoid copepods and diatoms. Virtually all prey items ingested by sardine were <1.2 mm maximum dimension, the particle size below which sardine only filter-feed. Despite the numerical dominance by phytoplankton, zooplankton contributed the major portion to sardine dietary carbon, small calanoid and cyclopoid copepods, anchovy eggs and crustacean eggs being the primary prey types. These results indicate that, like anchovy Engraulis capensis, sardine in the southern Benguela are primarily zoophagous, and contrast with earlier dietary studies on sardine in the region. However, the two species appear to partition their prey on the basis of size; sardine consume small zooplankton, whereas anchovy consume large zooplankton. This difference has been observed in other upwelling ecosystems where the two genera co-exist and is likely to contribute to the regime shifts observed between sardine and anchovy.     

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.     

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.     

The development and decline of phytoplankton blooms in the southern Benguela upwelling system. 2. Nutrient relationships

Five drogue studies on phytoplankton bloom dynamics in nutrient-rich newly upwelled water in the southern Benguela provided an opportunity to investigate the applicability of Redfield ratios to coastal upwelling regions. Mean atomic ratios of nutrient utilization and oxygen production (ΔP:ΔN:ΔSi:ΔO), as estimated from regression analyses (1:15,9:11,6:?283) and elemental changes along the drogue tracks (1:15,7:13,8:?206), were roughly similar to the equivalent Redfield ratios (1:16:15:?276). This implies that phytoplankton in upwelling areas also take up (or produce) these elements in proportions similar to those in which they occur in the cells themselves. As the upwelled water ages, changes in the ratios of N, Si and P imply that nitrate was used up fastest and is thus most likely to limit phytoplankton productivity in the southern Benguela region. However, regression analyses, calculations of times of nutrient depletion and published information on relative rates of nutrient regeneration suggest that silicate may also sometimes limit photosynthesis in this region.     

Influence of oceanographic fronts and low oxygen on the distribution of ichthyoplankton in the Benguela and southern Angola currents

The study focuses on ichthyoplankton populations in the southern Angola Current, the Angola-Benguela Front and the coastal upwelling area of the northern and central Benguela Current. The horizontal and vertical distributions of eggs and larvae of sardine Sardinops sagax, anchovy Engraulis encrasicolus and horse mackerel Trachurus trachurus capensis were examined in relation to distribution patterns of temperature, salinity and dissolved oxygen. Samples were collected during February–March 2002 using a Bongo net and a multiple opening/closing net in five depth layers (0–30m, 30–60m, 60–100m, 100–140m and 140–220m). Eggs and larvae were restricted mainly to the shelf areas in the Angola-Benguela Font and southern Angola Current waters. Their vertical distributions in the Angola-Benguela Front and northern Benguela upwelling region showed a sharp decline in abundance at about 60m depth. This decline appeared to be correlated to the distribution of dissolved oxygen in the water column It is hypothesised that low oxygen concentrations have a strong impact on the development and survival of the early life-cycle stages of fish, and that recruitment of sardine and other pelagic species relies more on the upward extension of the oxygen minimum layer than was previously thought.     

An individual-based model study of anchovy early life history in the northern Humboldt Current system

We used an individual-based model of anchovy (Engraulis ringens) early life history coupled with hydrodynamic outputs from the regional oceanic modeling system (ROMS) to investigate the factors driving variability in egg and larval survival rates in the northern Humboldt upwelling region off Peru. Individuals were released within a coastal area and followed for a period of 30 days. Those that were still in the coastal area at that time were considered as retained. We investigated the spatial and temporal variability in the release locations of the individuals retained, and compared these to observed egg concentration patterns reconstructed from a 40-year period of monitoring. A first set of simulations using passive particles to represent anchovy eggs and larvae revealed a large sensitivity of the results to the initial vertical distribution of particles. We then conducted two additional sets of simulations that included the effect of egg buoyancy, larval vertical swimming behavior and lethal temperature. We obtained (1) maximal coastal retention close to the surface in winter and in deeper layers in summer, (2) a large influence of egg buoyancy and of larval vertical behavior on coastal retention in all seasons, (3) a partial match between dates and locations of enhanced retention and observed egg concentration patterns and (4) a low effect of lethal temperature on survival except when associated with high egg density. The model suggests that an optimal temporal spawning pattern for maximizing coastal retention would have two maximums, the most significant in austral winter and the second in summer. This pattern agrees roughly with observed spawning seasonality, but with temporal discrepancy of about two months in the peaks of both series. Spatially, we obtained higher retention from 10 S to 20 S, whereas the observed maximum egg concentration was located between 6°S and 14°S. Among the three sets of simulations, the one taking into account larval vertical swimming behavior lead to the best match with the data.     

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.     

Simulated carbon and nitrogen flows of the planktonic food web during an upwelling relaxation period in St Helena Bay (southern Benguela ecosystem)

A vertically resolved ecosystem model is developed to simulate the dynamics of the pelagic food web in St Helena Bay during a representative period of relaxation after an upwelling event. The proposed model aims at coupling three biogeochemical cycles (carbon, nitrogen and silicon), using several recently developed concepts of the stoichiometric approach. A consequence of this approach is that important qualitative aspects are introduced, such as indicators of phytoplankton physiological state or variable food C:N ratios. For instance, the sedimentation and exudation rates for phytoplankton vary according to physiological state. An attempt is made to parameterize and simulate the diel cycles for vertical migration and feeding rhythms of large zooplankton, two important mesoscale processes that are thought to influence the overall dynamics of the huge phytoplankton blooms in the region.Observations of the Anchor Station Experiment 1987 (ASE’87) are used to assess the quality of the model. There is overall agreement between observations and the corresponding simulated results. The timing, the magnitude, and the vertical structure of the phytoplankton bloom are well reproduced. The balances for carbon and nitrogen flows and stocks compare well to the numerous estimates found from the literature for the southern Benguela region.On the basis of the model results, the origin of the new nutrients, the fate of the carbon fixed by phytoplankton, and the importance of the microheterotrophic pathways are discussed. It is concluded that sediments of the St Helena Bay and surrounding areas may play a crucial role in increasing the level of phytoplankton production. The results also suggest that exudation is the main process by which the carbon fixed by phytoplankton would have been lost, and that microheterotrophic pathways would have been intense during the experiment.     

The late Miocene onset of high productivity in the Benguela Current upwelling system as part of a global pattern

We have examined the history of the elevated primary productivity associated with the Benguela Current upwelling system off southwest Africa using sediments from 7.5 to 4.8 Ma at Ocean Drilling Program Site 1085 in the middle Cape Basin. Sedimentation rates are low until 6.9 Ma. Low accumulation rates of benthic foraminifers and organic carbon indicate that biological productivity was also low. Paleoproductivity dramatically increased at 6.7–6.5 Ma and was highly variable until 4.8 Ma with productivity maxima during cooler periods. The presence of radiolarian opal only between 5.8 and 5.2 Ma suggests an interlude of silica-rich intermediate water in the Cape Basin. The onset of heightened productivity under the Benguela Current is mirrored by similar increases reported between 6.9 and 6.7 Ma in the tropical eastern Pacific, the western and northern Pacific, and the Indian Ocean. The similarity between the patterns at Site 1085 and in the Pacific and Indian Oceans suggests that the dramatic productivity increase off southwest Africa is part of a global response to paleoceanographic changes.     

Phosphorus and nitrogen cyclings in the pelagic system of hiroshima bay: Results of numerical model simulation

A numerical ecosystem model expressing both phosphorus and nitrogen cyclings in the pelagic system of Hiroshima Bay, Japan, was developed in order to investigate the seasonal variations of these elements and their annual budgets. Based on the geophysical and chemical structures of the bay, Hiroshima Bay was divided into northern and southern boxes, themselves divided into two layers of an upper 5 m and a lower layer according to the stratification of the water column. The model consists of equations representing all relevant physical and biological processes. The results revealed that the internal regeneration of materials was an important source of bioavailable nutrients for phytoplankton growth in the water column. The incorporation of phytoplankton aggregation improved the accuracy of the outputs in comparison to the observed data, especially during the stratified summer season. The results also indicated that phosphorus limitation of phytoplankton growth occurs in the upper layer during summer while light limitation occurs in the lower layer. In addition, physical processes such as diffusion and advection were also important as they ensured that most of regenerated nutrients in the lower layer were transported to the upper layer. Thus, these processes might support the high primary production and the production of oysters that are extensively cultured in this bay. Considering the informative results obtained, the model used in this study provides a sound basis and tool to describe the dynamics of phosphorus and nitrogen cyclings in Hiroshima Bay.     

Vertical migration,catchability and acoustic assessment of semi-pelagic Cape horse mackerel Trachurus trachurus capensis in the southern Benguela

The vertical distribution and migration of the Cape horse mackerel Trachurus trachurus capensis were investigated on the basis of midwater and bottom trawl collections taken over diel cycles at two sites on South Africa's south coast. Attempts were also made to investigate the relationship between bottom trawl catch rates and acoustic abundance estimates. During the first cycle, horse mackerel were dispersed in midwater during the night, and towards sunrise they coalesced into schools and migrated to the seabed where they remained throughout the day. During that cycle, horse mackerel did not appear to feed, despite the prevalence of potential food, mainly Calanus copepods and euphausiid larvae, in the water column. In contrast, during the second cycle, horse mackerel remained near the seabed during the 24-h experiment, but they exhibited a marked feeding periodicity, with most feeding taking place in the late afternoon. They appeared to select for larger copepods in the water column. Several reasons are advanced to explain the difference in the vertical migratory behaviour of horse mackerel between the two sites, on the basis of the vertical distribution of their potential prey. Diel differences in acoustic estimates of horse mackerel abundance were significantly reduced after applying an acoustic deadzone correction factor. Trawl catches, however, could not be reconciled with acoustic estimates during the day, when horse mackerel were too close to the bottom for effective echo-integration.     

Studies on the role of marine bacteria in the upwelling ecosystem——Ⅰ.Bacterial biomass in the upwelling regions

-Six cruises were carried out off the south bank of Fujian - Taiwan during the period of December 21, 1987 to November 15, 1988 to estimate the contribution of bacterial biomass carbon (BBC) to the totai particulate organic pools using epifluorescent microscopic technique. The results show that the standing crop of bacteria fluctuated from 0. 95 to 66. 60 mg /m3 (dry weight). Upwelling phenomena appeared in the region around Nanpeng Island in summer while in the region of Waixie in all seasons. The average value of BBC was 27. 60(±6. 08)mg/m3and 21. 32 (±2. 34) mg/m3 respectively. The seasonal and spatial distribution is discussed in relation to environmental factors as well as upwelling phenomena. The role of bacteria in the flow of material and energy in the upwelling ecosystem is emphasized.