Northern Benguela Upwelling System Research Articles

Influence of oxygen minimum zone on macrobenthic community structure in the northern Benguela Upwelling System: a macro-nematode perspective

Influence of oxygen minimum zone on macrobenthic community structure in the northern Benguela Upwelling System: a macro-nematode perspective

Possible transport pathway of diazotrophic Trichodesmium by Agulhas Leakage from the Indian into the Atlantic Ocean.

Diazotrophic cyanobacteria such as Trichodesmium play a crucial role in the nitrogen budget of the oceans due to their capability to bind atmospheric nitrogen. Little is known about their interoceanic transport pathways and their distribution in upwelling regions. Trichodesmium has been detected using a Video Plankton Recorder (VPR) mounted on a remotely operated towed vehicle (TRIAXUS) in the southern and northern Benguela Upwelling System (BUS) in austral autumn, Feb/Mar 2019. The TRIAXUS, equipped with a CTD as well as fluorescence and nitrogen sensors, was towed at a speed of 8 kn on two onshore-offshore transects undulating between 5 and 200m over distances of 249km and 372km, respectively. Trichodesmium was not detected near the coast in areas of freshly upwelled waters but was found in higher abundances offshore on both transects, mainly in subsurface water layers down to 80m depth with elevated salinities. These salinity lenses can be related to northward moving eddies that most probably have been detached from the warm and salty Agulhas Current. Testing for interaction and species-habitat associations of Trichodesmium colonies with salinity yielded significant results, indicating that Trichodesmium may be transported with Agulhas Rings from the Indian Ocean into the Atlantic Ocean.

Low hypoxia tolerance in larvae of the sardine Sardinops sagax and anchovy Engraulis encrasicolus may limit their stock recovery in the northern Benguela upwelling system

Physiological traits of five key fish species (Cape hake Merluccius spp., pelagic goby Sufflogobius bibarbatus, Cape horse mackerel Trachurus capensis, European anchovy Engraulis encrasicolus and sardine Sardinops sagax) from the northern Benguela upwelling system (NBUS) were compared during their larval stage by measurements of enzymatic activities of key metabolic enzymes (citrate synthase [CS] and pyruvate kinase [PK]). Two distinct age classes (early larvae: 8‒14 days old; late larvae: 22‒29 days old) for each species and from two areas were compared: Terrace Bay (20° S), the main spawning ground of Cape horse mackerel and anchovy, and Kunene (17° S), near the Angola–Benguela frontal zone, where warm and hypoxic water masses intrude into the NBUS. The results revealed significantly higher CS activity in both larval age classes in Cape horse mackerel, anchovy and sardine compared with Cape hake and pelagic goby. Pelagic goby and Cape horse mackerel had significantly higher PK activity compared with Cape hake, anchovy and sardine, apparent in both larval age classes and both areas. Results for anaerobic metabolism indicate higher capacity for pelagic goby and Cape horse mackerel to recover from oxygen debt built up in prey capture and predator escape behaviour and a higher potential for hypoxia tolerance when compared with Cape hake, anchovy and sardine. These results suggest higher survival probability for pelagic goby and Cape horse mackerel compared with the other species under conditions of a shoaling oxygen minimum zone and may explain their current dominance in the NBUS.

Oxygen Variability in the Offshore Northern Benguela Upwelling System From Glider Data.

Despite their role in modulating the marine ecosystem, variability and drivers of low-oxygen events in the offshore northern Benguela Upwelling System (BenUS) have been rarely investigated due to the events' episodicity which is difficult to resolve using shipboard measurements. We address this issue using 4months of high-resolution glider data collected between February and June 2018, 100km offshore at 18°S. We find that oxygen (O2) concentrations in the offshore northern Benguela are determined by the subsurface alternation of low-oxygen Angola-derived water and oxygenated water from the south at 100-500m depth. We observe intermittent hypoxia (O2<60μmolkg-1) which occurs on average for ∼30% of the 4months deployment and is driven by the time-varying subsurface pulses of Angola-derived tropical water. Hypoxic events are rather persistent at depths of 300-450m, while they are more sporadic and have weekly duration at shallower depths (100-300m). We find extreme values of hypoxia, with O2 minima of 16μmolkg-1, associated with an anticyclonic eddy spinning from the undercurrent flowing on the BenUS shelf and showing no surface signature. Fine-scale patchiness and water mass mixing are associated with cross-frontal stirring by a large anticyclone recirculating tropical water into the northern BenUS. The dominance of physical drivers and their high variability on short time scales reveal a dynamic coupling between Angola and Benguela, calling for long-term and high-resolution measurements and studies focusing on future changes of both tropical O2 minima and lateral fluxes in this region.

High Diazotrophic Diversity but Low N2 Fixation Activity in the Northern Benguela Upwelling System Confirming the Enigma of Nitrogen Fixation in Oxygen Minimum Zone Waters

Oxygen minimum zones (OMZs) have been suggested as a suitable niche for the oxygen-sensitive process of biological fixation of dinitrogen (N2) gas. However, most N2 fixation rates reported from such waters are low. This low N2 fixation activity has been proposed to result from the unusual community of N2 fixers, in which cyanobacteria were typically underrepresented. The Northern Benguela Upwelling System (North BUS) is part of one of the most productive marine ecosystems and hosts a well-developed OMZ. Although previous observations indicated low to absent N2 fixation rates, the community composition of diazotrophs needed to understand the North BUS has not been described. Here, we present a first detailed analysis of the diazotrophic diversity in the North BUS OMZ and the Angola tropical zone (ATZ), based on genetic data and isotope speciation. Consistent with a previous study, we detected a slight N deficit in the OMZ, but isotope data did not indicate any active or past N2 fixation. The diazotroph community in the North BUS was dominated by non-cyanobacterial microbes clustering with members of gamma-proteobacteria, as is typical for other OMZ regions. However, we found a strikingly high diversity of Cluster III diazotrophs not yet described in other OMZs. In contrast to previous observations, we could also identify cyanobacteria of the clades Trichodesmium sp., UCYN-A and Cyanothece sp., in surface waters connected to or above the OMZ, which were potentially active as shown by the presence of genes and transcripts of the key functional marker gene for N2 fixation, nifH. While the detection of diazotrophs and the absence of active N2 fixation (based on isotopic speciation) are consistent with other OMZ observations, the detected regional variation in the diversity and presence of cyanobacteria indicate that we still are far from understanding the role of diazotrophs in OMZs, which, however, is relevant for understanding the N cycle in OMZ waters, as well for predicting the future development of OMZ biogeochemistry in a changing ocean.

Predatory species left stranded following the collapse of the sardine Sardinops sagax (Pappe, 1854) stock off the northern Benguela upwelling system: A review

Sardine Sardinops sagax (Pappe, 1854) is an important forage species, in the northern Benguela Upwelling System (nBUS). Sardine was a key forage species of the pelagic component in the nBUS, but the population collapsed due to a combination of overfishing and ecosystem change and variability in the nBUS. Multiple predators depended on sardine as high-quality food at the time when it was abundant. This study reviewed the current knowledge of predators' diet pre- and post-collapse of the sardine population stock in the nBUS. Data were sourced from previous studies on the trophic interaction in the nBUS. The historical and contemporary diets of commercially exploited species that prey on sardine were found to be relatively well documented while there are data deficiencies for non-commercially exploited species including cetaceans, sharks and some non-commercially exploited fish species. Amongst the species most reliant on sardine as prey are African penguin Spheniscus demersus, Cape gannet Morus capensis, Cape cormorant Phalacrocorax capensis, Cape monkfish Lophius vomerinus, shallow-water hake Merluccius capensis, deep-water hake M. paradoxus, snoek Thyrsites atun, Albacore tuna Thunnus alalunga, Cape fur seal Arctocephalus pusillus, Common dolphins Delphinus delphis, Dusky dolphins Lagenorhynchus obscurus, Heaviside's dolphins Cephalorhynchus heavisidii, six gills shark Hexanchus griseus, copper sharks Carcharhinus brachyurus, school shark Galeorhinus galeus and pelagic sharks Prionace glauca. Some predatory species such as the African penguin, Cape cormorant, Cape gannet, and school shark, are considered endangered as per the International Union for the Conservation of Nature (IUCN) Red List criteria. Populations of Albacore tuna and Cape monkfish are near threatened, while Cape fur seals, common dolphins, and shallow-water hake are considered the least concerned on the IUCN list. Although the collapse of sardine has affected the food webs it has not resulted in collapse, rather there have been changes. Three prey species; bearded goby, Cape horse mackerel, and jellyfish, were identified as alternative prey for some of the predators that previously fed on sardine. This review showed how overexploitation of sardine could have had far-reaching effects on the nBUS through feeding linkages. Goby, although a low-quality prey, is a successful substitute for sardine probably mostly due to its high abundance, tolerance to low oxygen and ability to thrive in warm waters. However, further studies are needed to establish the strength of the ecological linkages of the prey species that replaced sardine in the diets of the predators.

Oxygen and Nutrient Trapping in the Southern Benguela Upwelling System

The Benguela Upwelling System in the southeast Atlantic Ocean is of crucial socio-economic importance due to its high productivity. However, predicting its response to global change and understanding past changes are still great challenges. Here, we compile data obtained from a research cruise and an oceanographic mooring to demonstrate that a topographically steered nutrient trapping zone develops in a narrow belt along the coast during the main upwelling season in austral spring and summer in the southern Benguela Upwelling System. High nutrient concentrations within this zone increase the impact of upwelling on the productivity of the southern Benguela Upwelling System, but the efficient nutrient trapping operates at the expense of decreasing oxygen concentrations. This enhances the probability of anoxic events emerging toward the end of the upwelling season. However, at the end of the upwelling season, the front that separates the coastally trapped waters from open shelf waters weakens or even collapses due to upwelling cessation and the reversing current regime. This, in addition to a stronger vertical mixing caused by winter cooling, fosters the ventilation of the nutrient trapping zone, which reestablishes during the following upwelling season. The postulated intensification of upwelling and changes in the ecosystem structure in response to global warming seem to reduce the nutrient trapping efficiency by increasing offshore advection of surface waters and plankton blooms. The intensified upwelling and resulting lower biological oxygen consumption appears to mask the expected impacts of global warming on the oxygen minimum zone (OMZ) in the southern Benguela Upwelling System. In contrast to other OMZs, including those in northern Benguela Upwelling Systems, the OMZ in the southern Benguela Upwelling System reveals so far no detectable long-term decrease in oxygen. Thus, the nutrient trapping efficiency seems to be a critical feature mitigating global change impacts on the southern Benguela Upwelling System. Since it is topographically steered, regional impacts on the nutrient trapping efficiency appear also to explain varying responses of upwelling systems to global change as the comparison between southern and northern Benguela Upwelling System shows. This emphasizes the need for further and more comparable studies in order to better understand the response of Eastern Boundary Upwelling Systems and their ecosystem services to global change.

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

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

Coastal Trapped Wave Propagation along the Southwest African Shelf as Revealed by Moored Observations

AbstractCoastal trapped waves (CTWs) that propagate poleward along the southwest African shelf potentially leak energy from lower latitudes into the Benguela Upwelling System (BUS). Thus, in addition to local winds, these waves provide an important remote forcing mechanism for the upwelling region. The present study aims at elucidating the nature of CTWs in the northern BUS. To this end, we make use of multisite velocity observations from the Namibian shelf (18°, 20°, 23°S) and examine the alongshore velocity signal for signatures of CTWs by means of wavelet methods. We found that a substantial amount of energy is concentrated within a submonthly to subseasonal frequency band (10–50 days). Based on the coherence and phase spectra of the alongshelf currents, we provide evidence for a predominantly southward phase propagation and establish typical time and length scales of CTWs in the region. It turns out that their properties differ significantly within a few hundred kilometers along the coast. A comparison of the results with theoretical dispersion curves shows that this difference may be explained by variations in the bottom topography. Finally, we investigate the coupling of the alongshore currents with the coastal and equatorial wind stress and highlight regions of potential wave generation.

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

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

Pelagic key species and mechanisms driving energy flows in the northern Benguela upwelling ecosystem and their feedback into biogeochemical cycles

Abstract The northern Benguela Upwelling System (nBUS) has been facing increasing temperatures and decreasing dissolved oxygen (DO) levels over the last decades. This has implications for key processes and trophic interactions within the ecosystem including shifts in community composition, distribution ranges, and trophic levels, changes in energy flows and migration patterns with feedbacks to biogeochemical processes. Here we summarise the results gained from the GENUS project (Geochemistry and Ecology of the Namibian Upwelling System) focussing on the geochemical and ecological structures and processes dominating the pelagic component of the nBUS. Spatial and temporal distribution patterns of key species of zooplankton and fish larvae yielded biomass estimates (5 to 81 g Wet Mass m−2 (10 to 90% quantile) with a median of 19.5 g Wet Mass m−2 for the upper 200 m) and potential impacts on the vertical carbon flux. Vertical distribution ranges of key taxa were determined reflecting their specific abilities to tolerate hypoxia and, hence, their different adaptive mechanisms to cope with the Oxygen Minimum Zone (OMZ). The shoaling of the 2.5 mL O2 L−1-oxycline (0.24 m y−1) constrains sensitive species and hampers daily and seasonal vertical migrations. It may also affect the ability of organisms to maintain themselves within nearshore habitats by hindering vertical migration into deeper onshore currents. Respiration rates of key species were determined with one standard method (optode respirometry), showing an average respiration rate of 54.6 mL O2 d−1 (g Dry Mass)−1 for the bulk fraction of mesozooplankton, allowing also the estimate of DO consumption by mesozooplankton at different depth layers. Stable isotopic ratios (N, C) revealed trophic interactions and positions of zooplankton and fish. Our results reveal many players within a small range of trophic levels and a dominance of zooplankton taxa (copepods, euphausiids) in terms of biomass over small pelagic fish (sardine, anchovy), essential to consider for future higher-resolution ecosystem modelling.

Biogeochemical processes and turnover rates in the Northern Benguela Upwelling System

Abstract Biogeochemical cycles of carbon, nutrients, and oxygen transmit mean states, trends and variations of the physical realm in coastal upwelling systems to their food webs and determine their role in regional budgets of greenhouse gases. This contribution focuses on biogeochemical processes in the northern Benguela Upwelling System (NBUS), where low oxygen levels in upwelling source water are a major influence on carbon and nutrient cycles. Based on measurements during numerous expeditions and results of 3-D regional ecosystem modeling (project GENUS; Geochemistry and Ecology of the Namibian Upwelling System) we here examine source water character, effects of low oxygen conditions on nutrient masses and ratios, and of diazotrophic N2-fixation on productivity of the system and its transition to the adjacent eastern South Atlantic. In available observations, the effects of denitrification in water and sediment and phosphate release from sediments are minor influences on nitrate:phosphate ratios of the system, and excess phosphate in aged upwelling water is inherited from upwelling source water. Contrary to expectation and model results, the low N:P ratios do not trigger diazotrophic N2-fixation in the fringes of the upwelling system, possibly due to a lack of seeding populations of Trichodesmium. We also examine the flux of carbon from the sea surface to either sediment, the adjacent sub-thermocline ocean, or to regenerated nutrients and CO2. Observed fluxes out of the surface mixed layer are significantly below modeled fluxes, and suggest that regeneration of nutrients and CO2 is unusually intense in the mixed layer. This contributes to very high fluxes of CO2 from the ocean to the regional atmosphere, which is not compensated for by N2-fixation. Based on observations, the NBUS thus is a significant net CO2 source (estimated at 14.8 Tg C a− 1), whereas the CO2 balance is closed by N2-fixation in the model. Methane concentrations were low in surface waters in on-line measurements during 1 expedition, and based on these our estimate for the emission of methane for the entire Benguela system is below 0.2 Tg CH4 a− 1.

Upwelling indices for comparative ecosystem studies: Variability in the Benguela Upwelling System

Abstract The influence of climate variability on Eastern Boundary Upwelling Ecosystems (EBUEs) is evident through changes in productivity and shifts in species' distributions, yet to date, metrics of upwelling variability appropriate for comparative ecosystem studies have yet to be implemented. Here, we present synoptic-scale upwelling indices to quantify inter-annual to decadal variations in Ekman transport, at temporal and spatial scales relevant to the biota of EBUEs, and apply them to the Benguela Upwelling System (BUS). From 1979 to 2015, interannual, decadal-scale, and unidirectional variability in upwelling was observed, including a significant recent decrease in upwelling in the northern BUS, and a significant increase on the Agulhas Bank. These trends are associated with changes in the number of upwelling days and events in these regions, and correspond to a shift in the meridional positioning of the South Atlantic High pressure system.

Ecological relevance of salps and doliolids in the northern Benguela Upwelling System

Ecological relevance of salps and doliolids in the northern Benguela Upwelling System

Oxygen cycling in the northern Benguela Upwelling System: Modelling oxygen sources and sinks

This paper elucidates the oxygen dynamics in the northern Benguela Upwelling System by means of process oriented, numerical modelling. Owing to the complex physical-biological interaction in this system, a coupled hydrodynamic-biogeochemical model is required to grasp the various aspects of the oxygen dynamics. We used high-resolution atmospheric fields derived from observations to force our model, available since 1999. The model results represent a 15years, consistent data set of realistic hydrographic and ecosystem variables, including oxygen distribution patterns. After a concise description of the main aspects of the model, we use the model data to analyse the components contributing to the oxygen dynamics, namely, the ocean circulation, the exchange between ocean and atmosphere as well as the local biogeochemical oxygen cycling in the system. We thoroughly validate the model with available field observations and remote sensing data. The strengths of coastal upwelling, which controls the nutrient supply to the euphotic zone, as well as the poleward undercurrent that carries oxygen and nutrients to the shelf in the northern Benguela Upwelling System are well reproduced in the model. Among the biological oxygen sinks, mineralisation in the sediment, respiration of zooplankton and nitrification in the water column are important. We also found that vertical migration of zooplankton in response to the oxygen conditions provides a regulating feedback, which may prevent a complete deoxygenation of suboxic waters. As long as oxygen or nitrate are available in the bottom waters, the activities of chemolithoautotrophic sulphur bacteria on the sediment surface keep the redoxcline within the sediment and prevent the release of hydrogen sulphide into the water column. By horizontal integration of the simulated ocean-atmosphere oxygen flux, it can be shown that the Kunene upwelling cell between 16°S and 18°S is a boundary between the equatorial ocean, characterise by weak oxygen release to the atmosphere, and the subtropical Benguela Upwelling System governed by an enhanced and seasonal varying flux. Furthermore, a comparison of oxygen fluxes controlled by physical transport versus biogeochemical processes shows that the physical processes dominate in the northern Benguela Upwelling System.

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.

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.

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.

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.