Northern Benguela Research Articles

Measuring the recovery of the Northern Benguela Current Large Marine Ecosystem (BCLME): An application of the DPSIR framework

The objective of this paper is to describe the use of the Drivers, Pressures, State, Impact and Response (DPSIR) framework as a method for incorporating a multisector and multispecies approach for assessing progress in the recovery of a degraded Large Marine Ecosystem (LME). The Benguela Current Large Marine Ecosystem (BCLME) was degraded due to overfishing between 1960 and 1990. Various programmes were implemented to help the ecosystem to recover. These programmes included scientific studies, mechanisms to control exploitation of living marine resources, governance arrangements and capacity building. Progress made from these efforts was assessed using 3-year-moving average analyses and regression analyses of indicator trends. Twenty-seven indicators were analysed. The indicators chosen were based on the DPSIR framework and the modules of the LME approach. From the results, 41% of the indicators showed improvement of the ecosystem compared to 26% that showed continued degradation whilst the other 33% were neutral. Key improvement trends were seen in the slope values of indicators such as by-catch reduction 2566.42, total allowable catch (TAC) 7.63 and size abundances 0.60. Negative trends included biomass −170.24, and contribution of fisheries to GDP -0.35. Therefore, there were some negative effects but the BCLME is starting to show some signs of recovery. The results support the use of the LME approach as a sound way of addressing degraded marine ecosystems. The study also showed that using the DPSIR framework a multispecies, multidisciplinary approach can be achieved leading to a holistic review of the ecosystem.

Missing nitrogen fixation in the Benguela region

Opposing opinions on the importance of nitrogen fixation in the northern Benguela upwelling region provoked us to investigate the magnitude of nitrogen fixation in front of northern Namibia and southern Angola. Measurements of nitrogen fixation rates using the 15N method at 66 stations during seven cruises from 2008 to 2014 showed that, in general, the 15N content in the biomass did not increase after tracer incubation with 15N2, indicating that no nitrogen fixation occurred. Correspondingly, the filamentous nitrogen-fixing cyanobacterium Trichodesmium was almost not present. The abundant picocyanobacteria did obviously not perform nitrogen fixation to a significant degree. The artificial improvement of conditions for nitrogen fixation in mesocosm experiments, including phosphate and iron additions and a warmer temperature, failed to induce nitrogen fixation. A plausible explanation of these findings is a lack of conditioned cells for nitrogen fixation in the Benguela region.

The Principles of Buoyancy in Marine Fish Eggs and Their Vertical Distributions across the World Oceans.

Buoyancy acting on plankton, i.e. the difference in specific gravity between plankton and the ambient water, is a function of salinity and temperature. From specific gravity measurements of marine fish eggs salinity appears to be the only determinant of the buoyancy indicating that the thermal expansions of the fish egg and the ambient seawater are equal. We analyze the mechanisms behind thermal expansion in fish eggs in order to determine to what extent it can be justified to neglect the effects of temperature on buoyancy. Our results confirm the earlier assumptions that salinity is the basic determinant on buoyancy in marine fish eggs that, in turn, influence the vertical distributions and, consequently, the dispersal of fish eggs from the spawning areas. Fish populations have adapted accordingly by producing egg specific gravities that tune the egg buoyancy to create specific vertical distributions for each local population. A wide variety of buoyancy adaptations are found among fish populations. The ambient physical conditions at the spawning sites form a basic constraint for adaptation. In coastal regions where salinity increases with depth, and where the major fraction of the fish stocks spawns, pelagic and mesopelagic egg distributions dominate. However, in the larger part of worlds’ oceans salinity decreases with depth resulting in different egg distributions. Here, the principles of vertical distributions of fish eggs in the world oceans are presented in an overarching framework presenting the basic differences between regions, mainly coastal, where salinity increases with depth and the major part of the world oceans where salinity decreases with depth. We show that under these latter conditions, steady-state vertical distribution of mesopelagic fish eggs cannot exist as it does in most coastal regions. In fact, a critical spawning depth must exist where spawning below this depth threshold results in eggs sinking out of the water column and become lost for recruitment to the population. An example of adaptation to such conditions is Cape hake spawning above the critical layer in the Northern Benguela upwelling ecosystem. The eggs rise slowly in the onshore subsurface current below the Ekman layer, hence being advected inshore where the hatched larvae concentrate with optimal feeding conditions.

Trophic relationships of hake (Merluccius capensis and M. paradoxus) and sharks (Centrophorus squamosus, Deania calcea and D. profundorum) in the Northern (Namibia) Benguela Current region

The trophic relationships of two hake species (Merluccius capensis and M. paradoxus) and three shark species (Centrophorus squamosus, Deania calcea and D. profundorum) were investigated using nitrogen and carbon stable isotope signatures (δ15N and δ13C) of their muscle tissues. The sharks were more enriched in 15N than the hake, an indication of the apex predator status of sharks. Among the sharks considered, C. squamosus occupied the highest trophic level and fed primarily on benthic prey. The two species of shark from the genus Deania were not different based on δ15N or δ13C, so they had similar diets. The δ13C signatures indicated that M. capensis and sharks fed on prey derived from similar basal resources. However, there was a significant difference in δ13C between M. paradoxus and all other species examined, suggesting that they occupied different feeding niches. Isotope-based population metrics showed narrower trophic ranges in sharks than M. capensis. Carbon and nitrogen ranges indicated that hake fed on a more diverse pool of carbon sources and had generally more enhanced trophic diversity in their feeding patterns than sharks. Among the species considered, C. squamosus occupied a unique isotopic space. Our results supported the hypothesis there are trophic overlaps among these species, but some interesting differentiation was revealed.

Spawning patterns of shallow-water hake (Merluccius capensis) and deep-water hake (M. paradoxus) in the Benguela Current Large Marine Ecosystem inferred from gonadosomatic indices

We use gonad- and body-weight data from 54,000 samples of Merluccius capensis and Merluccius paradoxus collected in all months of the years between 1991 and 2013 to infer peak spawning periods and areas in the Benguela Current Large Marine Ecosystem. We develop and apply a new gonadosomatic index threshold for identification of spawning individuals. Spawning M. capensis were observed throughout the study area, mainly in areas of about 100m bottom depth. The highest proportions of spawning M. capensis females in the northern Benguela region were observed off central Namibia between 24.0 and 26.0°S. In the southern Benguela, peaks in the proportions of spawning M. capensis were observed in two areas off the South African West Coast (31.0–32.5°S and 34.5–36.0°S), whereas spawning females off the South African South Coast (east of 20°E) appeared to be more evenly distributed in space. Seasonality differed between areas. In the northern Benguela, the main spawning season of M. capensis appeared to be the austral winter (July–September, peaking in August), while off the South African South Coast, the main spawning season is suggested to be in summer (around January). Between these two extremes, on the western Agulhas bank in the southern Benguela, spawning peaks were observed in both summer and winter. These peaks largely coincided with peaks in phytoplankton production that are linked to upwelling conditions in the region. Hake condition decreased subsequent to the development of the gonads. The annual spawning cycle differed between small and large M. capensis. The current October–closure of the fishery in Namibia may not match the peak spawning in August/September and may need to be shifted to earlier in the year. Spawning M. paradoxus were mainly found in areas of 200–650m bottom depths. In the northern Benguela, spawning M. paradoxus were observed as far north as 25°S in August. The proportion of spawning females peaked between 34.5°S and 36.5°S off the West Coast, and between 23.0°E and 26.5 °E off the South Coast. It was suggested that M. paradoxus spawn throughout the year off the South African coast, with increased intensity around March and August–October. The finding of multiple spawning seasons and areas of both M. capensis and M. paradoxus strongly suggest multiple stocks (reproductive units).

Life in the sea of plenty: Seasonal and regional comparison of physiological performance of Euphausia hanseni in the northern Benguela upwelling system

Variability in upwelling events may lead to periods of constrained food availability in the northern Benguela upwelling system (NBUS), thereby affecting the physiological state and metabolic activity of euphausiids. Most attention has so far been paid to seasonal effects but little is known about regional variability. Metabolic activity (expressed by respiration and excretion rates) and physiological state (expressed by reproductive effort and moult activity) in Euphausia hanseni were examined at different stations during austral summer (minimum upwelling) and austral winter (maximum upwelling). Overall, regional differences in physiological state, influencing metabolic activity, were greater than seasonal ones, indicating favourable conditions for growth and reproduction year-round. Higher respiration rates were found for females in more advanced stages of sexual development. Moult stage did not affect oxygen consumption rates, however. The physiological state of E. hanseni at the time of capture may serve as a meaningful indicator of the associated hydrographic conditions in the NBUS, to be further used in eco-system analysis on seasonal or long-term time scales. A latitudinal comparison of species highlights the extraordinary physiological plasticity of euphausiids.

Zonation of bacterioplankton communities along aging upwelled water in the northern Benguela upwelling.

Upwelling areas are shaped by enhanced primary production in surface waters, accompanied by a well-investigated planktonic succession. Although bacteria play an important role in biogeochemical cycles of upwelling systems, little is known about bacterial community composition and its development during upwelling events. The aim of this study was to investigate the succession of bacterial assemblages in aging upwelled water of the Benguela upwelling from coastal to offshore sites. Water from the upper mixed layer at 12 stations was sampled along two transects from the origin of the upwelling to a distance of 220 km. 16S rRNA gene amplicon sequencing was then used in a bacterial diversity analysis and major bacterial taxa were quantified by catalyzed reporter deposition-fluorescence in situ hybridization. Additionally, bacterial cell numbers and bacterial production were assessed. Community statistical analysis revealed a reproducible zonation along the two transects, with four clusters of significantly different microbial assemblages. Clustering was mainly driven by phytoplankton composition and abundance. Similar to the temporal succession that occurs during phytoplankton blooms in temperate coastal waters, operational taxonomic units (OTUs) affiliated with Bacteroidetes and Gammaproteobacteria were dominant during algal blooming whereas “Pelagibacterales” were highly abundant in regions with low algal abundance. The most dominant heterotrophic OTU (9% of all reads) was affiliated with “Pelagibacterales” and showed a strong negative correlation with phytoplankton. By contrast, the second most abundant heterotrophic OTU (6% of all reads) was affiliated with the phylum Verrucomicrobia and correlated positively with phytoplankton. Together with the close relation of bacterial production and phytoplankton abundance, our results showed that bacterial community dynamics is strongly driven by the development and composition of the phytoplankton community.

Biannual otolith‐zone formation of young shallow‐water hake Merluccius capensis in the northern Benguela: age verification using otoliths sampled by a top predator

Otoliths collected at least monthly from scat samples of Cape fur seals Arctocephalus pusillus are used to show that shallow-water hake Merluccius capensis from the northern Benguela develop three translucent zones in their first 1·5 years of life. The novel sampling approach provided otoliths that belonged to four M. capensis cohorts of approximate known age (hatched in 1996, 1998, 2002 and 2005), allowing age verification. Following spawning in austral winter, translucent zones consistently formed first in summer and autumn (T1), then in winter and spring (T2) and again in summer and autumn (T3), with no difference in appearance of the zones (biannuli) for the four cohorts considered. The second translucent zone is usually the first true annulus (year mark). It forms during July to September in fish of 15-20 cm total length (LT ). Formation of the translucent zones appears to be determined by fish length or age, rather than by exogenous cues. It is suggested that length measurements should be used to help determine the first age group; fish with a translucent zone marked at otolith lengths >7·5 mm should be termed 1 year-old fish. Ages of M. capensis used in previous stock assessment models have been overestimated. Biannuli are an unusual occurrence in fish otoliths in general, but have been observed in other Merluccius species.

Late Pliocene upwelling in the Southern Benguela region

The Late Pliocene has been proposed as a possible analogue for understanding future climate change and for testing climate models. Previous work has shown that during the Pliocene the major upwelling systems were relatively warm, and that this meant they were either inactive, contracted, or were upwelling warmer waters than present. Here, we examine evidence from a site located on the margins of the modern Benguela Upwelling system to test whether the upwelling cells had migrated or contracted relative to present during the Pliocene. We applied several organic geochemistry proxies and foraminiferal analyses to reconstruct the Pliocene history of ODP Site 1087 (31°28′S, 15°19′E, 1374 m water depth), including the UK37′ and TEX86 indices (for reconstructing sea surface temperatures), phytoplankton biomarker concentrations and stable isotope ratios (for estimating export primary productivity, and for oxygen isotope statigraphy), and planktonic foraminifera assemblage abundances (for inferring water mass changes). These proxies show that, between 3.5 and 3.0 Ma, the southern Benguela region was cooler than the northern Benguela region by 5 °C, the latter being where the main upwelling cells are found today. From the multiproxy data obtained, we also infer that more extensive upwelling was present in the southern Benguela region during the Pliocene than at present, and that the Benguela Upwelling cells shifted northwards after the Pliocene epoch as a result of changes in the local wind field. We also find evidence that the Benguela Upwelling was sensitive to the pronounced cooling during the M2 and KM2 glacial stages, potentially associated with the expansion of sea ice and cooling in Antarctica in the Late Pliocene.

Downscaling biogeochemistry in the Benguela eastern boundary current

Dynamical downscaling is developed to better predict the regional impact of global changes in the framework of scenarios. As an intermediary step towards this objective we used the Regional Ocean Modeling System (ROMS) to downscale a low resolution coupled atmosphere–ocean global circulation model (AOGCM; IPSL-CM4) for simulating the recent-past dynamics and biogeochemistry of the Benguela eastern boundary current. Both physical and biogeochemical improvements are discussed over the present climate scenario (1980–1999) under the light of downscaling.Despite biases introduced through boundary conditions (atmospheric and oceanic), the physical and biogeochemical processes in the Benguela Upwelling System (BUS) have been improved by the ROMS model, relative to the IPSL-CM4 simulation. Nevertheless, using coarse-resolution AOGCM daily atmospheric forcing interpolated on ROMS grids resulted in a shifted SST seasonality in the southern BUS, a deterioration of the northern Benguela region and a very shallow mixed layer depth over the whole regional domain. We then investigated the effect of wind downscaling on ROMS solution. Together with a finer resolution of dynamical processes and of bathymetric features (continental shelf and Walvis Ridge), wind downscaling allowed correction of the seasonality, the mixed layer depth, and provided a better circulation over the domain and substantial modifications of subsurface biogeochemical properties. It has also changed the structure of the lower trophic levels by shifting large offshore areas from autotrophic to heterotrophic regimes with potential important consequences on ecosystem functioning. The regional downscaling also improved the phytoplankton distribution and the southward extension of low oxygen waters in the Northern Benguela. It allowed simulating low oxygen events in the northern BUS and highlighted a potential upscaling effect related to the nitrogen irrigation from the productive BUS towards the tropical/subtropical South Atlantic basin. This study shows that forcing a downscaled ocean model with higher resolution winds than those issued from an AOGCM, results in improved representation of physical and biogeochemical processes.

Spawning and nursery areas, longitudinal and cross‐shelf migrations of the Merluccius capensis stock in the northern Benguela

AbstractWe investigate the spatial distributions of juvenile and adult hake Merluccius capensis on the Namibian continental shelf using data from 25 biomass surveys (1990–2007) to identify (i) nursery/spawning areas, their spatial or temporal separation and change and (ii) length, depth and latitude preference patterns. The density of fish (number‐of‐fish per 30‐min‐haul) was examined in relation to fish total length (TL) and latitude and TL and bottom depth. Nursery aggregations were most frequent in central (22–25°S) and southern (26–29°S) Namibia, increasing in density in the south since 2000. Hatch dates of ≤17 cm fish were calculated from juvenile growth rates. Peaks occurred in winter and summer‐autumn in the centre and slightly later in the south. Spawning areas appear to have shifted southward since the late 1970s, showing the plasticity of this stock as a response to fishing pressure and environmental variability and change. Merluccius capensis show a stable pattern of latitude preference over the 25 surveys examined. They first occur on the mid‐shelf <9 cm, generally moving to the inner‐shelf at 9–15 cm (<1 yr old). They generally prefer the northern and mid‐shelf area between 24 and 45 cm (1.5–3.5 yr old), probably for feeding and building resources for spawning purposes. They later move to the outer‐shelf and return southward and to the mid‐shelf region to spawn at ≥45 cm TL (≥3.5 years old), a contained stock unit in the northern Benguela. We propose a complete migration life history of M. capensis for the first time, showing their extensive longitudinal migrations, similar to Merluccius species elsewhere.

Synthesis: climate effects on biodiversity, abundance and distribution of marine organisms in the Benguela

AbstractThe NansClim project (2010–2013) represented a regional collaboration to assess the effects of climate on Benguela dynamics. Based on in situ (since the 1960s in Namibia and South Africa and 1985 in Angola) and satellite (since the 1980s) observations, the project focussed on four subsystems, namely the Angola subtropical, northern Benguela upwelling, southern Benguela upwelling and Agulhas Bank. This contribution summarizes the findings for selected key questions, ranging from changes in the physico‐chemical habitats, plankton, pelagic and demersal fish communities, to cross‐cutting evaluation at subsystem and regional scales. The results underline the overriding importance to of considering the combined effects of climate and fishing as drivers of the dynamics of the ecosystem components. Each subsystem currently continues to function largely as a separate entity as described in earlier reviews. However, some changes have been observed across several subsystems, e.g., a coherent shift from one relatively stable period to another occurred in the northern and southern Benguela in the mid‐1990s. Future climate change could weaken the boundaries between the four subystems. The findings underline the need for continued regional research collaboration and regional surveys focussed at ecosystem, rather than resource, assessment. Our conclusions include implications for ecosystem‐based fisheries management, and recommendations for future regional research.

The relation of wind stress curl and meridional transport in the Benguela upwelling system

Spatially high resolved wind data reveals the existence of a very pronounced and persistent cyclonic (negative) wind stress curl (WSC) along the southwest African coast. Several theoretical studies have shown that negative WSC modifies the meridional currents in eastern boundary systems significantly. Since the water mass composition on the southwest African shelf is primarily controlled by the meridional currents, understanding the relation between WSC and the meridional advection is of great importance for the Benguela ecosystem. In this study, we use a regional general circulation model that is validated with observations in order to study the effect of cyclonic WSC on the meridional transport along the southwest African coast. We show that there is a connection between the meridional transport and the WSC on a seasonal time scale in the northern Benguela upwelling system (BUS). The meridional transport follows the annual and semi-annual cycles of the WSC between 20° S and 25° S. The cyclonic WSC in the northern BUS may therefore support a southward advection of tropical waters into the upwelling system.

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

AbstractSpatial 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.

Distribution, feeding behaviour, and condition of Cape horse mackerel early life stages, Trachurus capensis, under different environmental conditions in the northern Benguela upwelling ecosystem

AbstractEarly life history (ELH) traits are key to understand variable recruitment success and hence the stock size of marine fish. One of the currently most puzzling ecosystems in this regard is the northern part of the Benguela Current upwelling system off Namibia. Here, populations of the formerly dominant pelagic species, sardine and anchovy, failed to recover during the last three decades after a dramatic decline. In contrast, Cape horse mackerel, Trachurus capensis, maintained a constant population size. Warming of the system and shoaling of hypoxic zones together with feedback loops within an altered foodweb are discussed to be responsible for this regime shift. In this study, we address the role of larval traits for the successful performance of the T. capensis population under the present environmental conditions with the focus on feeding ecology. We investigated seasonal variations of the geographical distribution, growth rate, feeding ecology, and nutritional condition of their ELH stages and examined relationships with water temperature, dissolved oxygen concentration, and micro-zooplankton composition. T. capensis' ELH stages showed a wide spatial and seasonal distribution, a preference for higher water temperatures (18–21°C) and presence over a wide range of dissolved oxygen concentrations (0.13–6.35 ml O2 l−1). Feeding success was high and mainly different groups of Copepoda were targeted, which were strongly size selected. The high dietary importance of micro-copepods during large parts of the larval phase indicates successful exploitation of this food source, which has increased in abundance during the last decade. It also explains observed best nutritional conditions at temperatures between 18 and 21°C, since these small copepods are commonly associated with warmer temperatures. Altogether, these traits enhance the species' probability to encounter suitable environments for the survival of their ELH stages, which is likely to lead to their high recruitment success in the northern Benguela ecosystem.

Distribution of zooplankton biomass and potential metabolic activities across the northern Benguela upwelling system

The distribution of zooplankton biomass and potential metabolic rates, in terms of electron transport system (ETS) and glutamate dehydrogenase (GDH), were analyzed along a cross-shelf transect in waters off Namibia. The highly variable dynamics of upwelling filaments promoted short-term fluctuations in the zooplankton biomass and metabolism. Maximum values were characteristically found over the shelf-break, where zooplankton biomass as dry mass (DM) reached peaks of 64.5mgm−3 within the upper 200m in late August. Two weeks later, the zooplankton-DM decreased by more than a third (19mg DM m−3). Zooplankton potential respiration and NH4+ excretion averaged 234μmol O2 m−3 d−1 and 169μmol NH4+m−3 d−1 in the Namibian shelf, respectively. High protein-specific ETS activities even in the low-chlorophyll waters outside the filament suggested a shift into greater omnivory seaward. In this light, zooplankton elemental and isotopic compositions were used to investigate the pelagic food web interactions. They evidenced spatial changes in the carbon resource for zooplankton as well as changes in the form of nitrogen that fueled the biological production in aging advected waters. Overall, both aspects of zooplankton metabolism impacted the primary productivity at a level less than 10% under all the different oceanographic conditions.

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.

Succession of micro- and nanoplankton groups in ageing upwelled waters off Namibia

The phytoplankton composition was investigated on a cross-shore transect in the Northern Benguela upwelling system in August and September 2011. Micro- and nanorganisms were microscopically analysed and assigned to appropriate phytoplankton groups. The distribution of diatoms, dinoflagellates, coccolithophores and nanoflagellates along the transect was studied in order to reflect the phytoplankton succession. Typical phytoplankton communities indicating different stages of succession were found in relation to the distance to the coast and in relation to the aged upwelled water. Three succession stages could be well defined and verified by satellite data in the newly upwelled water, matured water and aged offshore water. The different stages were distinguished not only by the significant differences in the nutrient concentrations but also by the dominating species and phytoplankton life forms.A mixed population of dinoflagellates, coccolithophores and microflagellates was observed in the newly upwelled water. Small diatom cells (Cylindrotheca closterium) marked the transition to the matured water at about 40km from the shore. A diatoms to dinoflagellates ratio up to 33 and biomasses up to 250μgCl−1 were characteristic for the matured stage, between 40km and 250km from the coast. As observed by satellite data, highly productive water bands intrude the water masses which also represent this stage. They were characterized by accumulated biomasses dominated by diatoms (Pseudo-nitzschia seriata group). Diatoms to dinoflagellates ratios less than 1 and maximum biomasses of 15μgCl−1 characterized the aged water at the later stage of succession. Abundances of dinoflagellates smaller than 10μm increased just after the decline of the diatoms.

Cross shelf hydrographic and hydrochemical conditions and their short term variability at the northern Benguela during a normal upwelling season

Cross shelf hydrographic and hydrochemical conditions were investigated during the seasonal maximum of upwelling in the northern Benguela upwelling system. The study combines in situ observations, remotely sensed data and results of a regional 3-dimensional numerical model. In situ observations were recorded along a cross shelf transect off Namibia starting at 20°S 13°E, repeated five times during 16 August 2011 until 19 September 2011. Comparison of wind forcing and sea surface temperatures during the time of the expedition with long-term climatological data as well as the index of intensity of the Benguela upwelling indicates “normal” upwelling conditions in austral winter 2011 in the northern Benguela. Small scale temporal (days) and spatial (km) variability is high during the upwelling season, primarily caused by highly variable wind forcing and dynamics of mesoscale structures like eddies and filaments as found in remotely sensed data. This mesoscale dynamics impact the applicability of a conceptual 2-dimensional circulation model, i.e. a linear succession along the cross-shelf transect. Therefore, an age proxy for surface water was constructed based on oxygen and heat fluxes during the first aging period and on salinity and heat fluxes during the second phase. The application of an age proxy instead of distance to shore successfully validates the succession concept. Furthermore, the investigation of the upwelling strengths by analytical and circulation models verified their dependence on coastal- and curl driven upwelling processes with the onshore dominance of coastal upwelling. In the investigated time period, offshore, curl driven upwelling dominated with a maximum located on the shelf.

Distribution and paleoecology of molluscan skeletal remains along an upwelling tract: Benguela system, Namibian shelf

Abstract This study describes and categorizes the preservable, and thus predominantly molluscan, component of the macrofauna death assemblages of sub-environments within the northern Benguela upwelling system. This is especially important for assembling actualistic data that can later serve for identifying similar settings in the geological record. A total of 16 samples were examined. Contrary to stereotypes, benthic communities are present but highly variable across the high-productivity tract and are characterized by distinctive community structures. Benguela system document a decrease in body size of chemosymbiotic bivalves (lucinids) and percent of chemosymbionts and deposit-feeders, and an overall increase in species richness and epifaunal-infaunal ratio. This trend is in accordance with the upwelling facies-gradient from diatomaceous ooze in areas of maximum upwelling intensity outward to the aerated but food-abundant carbonate oozes and spatially coexisting phosphorite concentrations. Differences in seafloor oxygen concentration along with variable food availability played major roles in structuring the death assemblages under the Benguela upwelling system. The ecologic data provide ways to distinguish between facies of low and high shell production along an increasing oxygen gradient, and also between assemblages with moderate and higher species richness associated with time-averaging. The data support the existence of a wider range of time averaging in the formation of molluscan sand and gravel samples in the northern Benguela than previously appreciated. Given the scarcity of information on macrobenthic ecology from high-productivity settings, these death assemblages can represent a modern analog.