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

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

Environmental influence on phytoplankton production during summer on the KwaZulu-Natal shelf of the Agulhas ecosystem

During February 2010, studies of primary production (PP) and physiology were conducted at five selected sites in the KwaZulu-Natal (KZN) Bight of the Agulhas ecosystem as part of a programme to elucidate the influence of major physical driving forces and nutrient inputs on the structure and functioning of biological communities. These sites were located in the vicinity of the Durban lee eddy, in the midshelf region of the central part of the bight, off the Thukela Mouth, and to the north and south of Richards Bay. At four of the sites, chlorophyll a ranged from 0.10 to 1.44?mg m–3 and integrated PP ranged between 0.35 and 2.58?g C m–² d–1. The highest biomass and PP, which were comparable to those observed in a wind-driven upwelling system, were associated with a diatom community observed at the midshelf site, and varied between 0.26 and 4.27?mg m–³ and 7.22 and 9.89?g C m–² d–1, respectively. Environmental conditions at each of the sites differed substantially and appeared to be influential in initiating and controlling the development and distribution of phytoplankton biomass and production. Phytoplankton adaptation to variable environmental conditions was characterised by a decreased light-limited slope (αB) and increased rate of photosynthesis (P_m ) and light saturation (Ek) with elevated temperatures. The converse (increased α^B and decreased P_m and Ek) was observed as irradiance levels declined. Generalised additive models indicated that irradiance, temperature and biomass were important variables influencing photosynthetic parameters and photosynthetic rates.     

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

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

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

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

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.     

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.     

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.     

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.     

Estimates of phytoplankton and bacterial biomass and production in the northern and southern Benguela ecosystems

Available data on phytoplankton and bacterial abundance and production off the coasts of southern Africa (to the 500 m depth contour) have been assembled and analysed for a network analysis of carbon flow in the Benguela ecosystem. Phytoplankton carbon biomass (from measurements of chlorophyll a) in the northern Benguela (2 558 300 tons) was considerably higher than in the southern Benguela (671 420 and 516 400 tons for the West and South coasts respectively). However, overall annual production (from C¹⁴-uptake measurements) was similar, 77 416 608, 76 399 973 and 78 988 020 tons C·year^?1 respectively. Phytoplankton respiration and sedimentation losses were calculated as functions of primary production and therefore followed similar trends. From the most conservative estimates (mean bacterial biomass of 10 mg C·m^?3 and average P:B of 0,2·day^?1) bacterial biomass is 2–7 per cent of phytoplankton biomass in the northern and southern Benguela, and bacterial production is 3–5 per cent of primary production. Assuming a net growth yield of 30 per cent, bacteria would need to consume 9–15 per cent of the total primary production in order to meet their requirements for carbon consumption. Calculations based on a mean bacterial biomass of 40 mg C·m^?3 and a mean growth rate of 0,5·day^?1 in the upper 30 m of the water column show bacterial biomass to be 8–27 per cent of phytoplankton biomass and bacterial production to be 26–44 per cent of phytoplankton production. Bacterial carbon consumption requirements at these rates amount to 86–147 per cent of total primary production.     

Size fractionated biomass and productivity of phytoplankton and new production in the Prydz Bay and the adjacent Indian sector of the Southern Ocean during the austral summer 1998/1999

INTRODUCTIONComparedwithotheroceansintheworld ,thereisrelativelyhigherlevelofnutrients (ni trate ,phosphate ,silicate)intheSouthernOcean .Butprimaryproductivitiesarelow ,andnewproductionisalsoonlyinthemiddlelevelthere .Itiscommonlyconsideredthatthelowerpro ductivitiesintheSouthernOceaniscausedbylowtemperature ,lowlight,lowstabilityofwa ter,scarcityoftraceelementsuchasironandgrazingbyzooplankton (Burkilletal.,1 995) ,etc.ThemostexistedstudiesconcentratedontheAtlanticSectoroftheSouthernO…     

南黄海浮游植物初级生产力粒级结构与碳流途径分析

根据2006年夏季与冬季南黄海真光层内浮游植物初级生产速率的实测数据,对比研究了两种差异显著的水动力条件下,不同粒级浮游植物初级生产力水平、时空分布特征及其环境调控机制,并探讨了真光层生源碳可能的碳流途径。研究结果表明,南黄海夏季总初级生产力(碳)平均为30.69 mg/(m2·h),高值区位于调查海域南部长江冲淡水影响区,冬季总初级生产力低于夏季,平均水平为21.73 mg/(m2·h),高值区北迁至海州湾附近;夏季不同粒径浮游植物对总初级生产的贡献率由高到低的顺序为小型(42.8%)、微型(29.6%)、微微型(27.6%),冬季的为微微型(41.2%)、微型(36.5%)、小型(22.3%);真光层初级生产力与环境因子的相关分析表明,水动力条件(混合与层化)引起的营养盐来源和光照的变化是初级生产力分布的主要控制因素;用生态比值法对南黄海碳流途径的分析表明,微食物环在冬夏两季碳流途径中均占有重要地位,从总体上看,南黄海浮游生态系统的碳输出潜力较弱。     

Nutrient controls on new production in the Bodega Bay, California, coastal upwelling plume

A theoretical framework for the time-dependent processes leading to the high rates of new production in eastern boundary upwelling systems has been assembled from a series of past upwelling studies. As part of the CoOP WEST (Wind Events and Shelf Transport) study, new production in the Bodega Bay upwelling area and it's control by ambient nitrate and ammonium concentrations and the advective wind regime are described. Data and analyses are focused primarily on the WEST 2001 cruise (May–June 2001) when the two legs differed greatly in wind regimes but not nutrient concentrations. Elevated concentrations of ammonium in upwelled water with high nitrate were observed in both legs. Nitrate uptake by phytoplankton as a function of nitrate concentration was linear rather than Michaelis–Menten-like, modulated by inhibitory levels of ammonium, yielding coefficients that enable the specific nitrate uptake element of new production to be estimated from nutrient concentrations. The range of specific nitrate uptake rates for the two legs of WEST 2001 were similar, essentially a physiological response to nutrient conditions. However, the low “realization” of new production i.e. incorporation of biomass as particulate nitrogen that occurred in this system compared to the theoretical maximum possible was determined by the strong advective and turbulent conditions that dominated the second leg of the WEST 2001 study. These data are compared with other upwelling areas using a physiological shift-up model [Dugdale, R.C., Wilkerson, F.P., Morel, A. 1990. Realization of new production in coastal upwelling areas: a means to compare relative performance. Limnology and Oceanography 35, 822–829].     

Modelling the effect of food availability on recruitment success of Cape anchovy ichthyoplankton in the southern Benguela upwelling system

Cape anchovy Engraulis encrasicolus adapted its reproductive strategies to the southern Benguela system by spawning over the Agulhas Bank, an area of low productivity that is located upstream of the predominant upwelling system. Frontal jet currents transport eggs and larvae toward the west coast of South Africa, where recruitment takes place. To characterise the recruitment dynamics of Cape anchovy ichthyoplankton, we used an individual-based model forced by a coupled hydrodynamic–biogeochemical model. The results show the importance of food (especially diatoms and copepods) dynamics on the spatial and temporal patterns of recruitment success, and also confirm the importance of the spawning area, timing and water depth on the recruitment success of Cape anchovy larvae.     

Effects of variable winds on biological productivity on continental shelves in coastal upwelling systems

The production and distribution of biological material in wind-driven coastal upwelling systems are of global importance, yet they remain poorly understood. Production is frequently presumed to be proportional to upwelling rate, yet high winds can lead to advective losses from continental shelves, where many species at higher trophic levels reside. An idealized mixed-layer conveyor (MLC) model of biological production from constant upwelling winds demonstrated previously that the amount of new production available to shelf species increased with upwelling at low winds, but declined at high winds [Botsford, L.W., Lawrence, C.A., Dever, E.P., Hastings, A., Largier, J., 2003. Wind strength and biological productivity in upwelling systems: an idealized study. Fisheries Oceanography 12, 245–259]. Here we analyze the response of this model to time-varying winds for parameter values and observed winds from the Wind Events and Shelf Transport (WEST) study region. We compare this response to the conventional view that the results of upwelling are proportional to upwelled volume. Most new production per volume upwelled available to shelf species occurs following rapid increases in shelf transit time due to decreases in wind (i.e. relaxations). However, on synoptic, event time-scales shelf production is positively correlated with upwelling rate. This is primarily due to the effect of synchronous periods of low values in these time series, paradoxically due to wind relaxations. On inter-annual time-scales, computing model production from wind forcing from 20 previous years shows that these synchronous periods of low values have little effect on correlations between upwelling and production. Comparison of model production from 20 years of wind data over a range of shelf widths shows that upwelling rate will predict biological production well only in locations where cross-shelf transit times are greater than the time required for phytoplankton or zooplankton production. For stronger mean winds (narrower shelves), annual production falls below the peak of constant wind prediction [Botsford et al., 2003. Wind strength and biological productivity in upwelling systems: an idealized study. Fisheries Oceanography 12, 245–259], then as winds increase further (shelves become narrower) production does not decline as steeply as the constant wind prediction.     

Photosynthate allocation in a temperate sea over an annual cycle: the relationship between protein synthesis and phytoplankton physiological state

The seasonal and vertical variations in the patterns of photosynthate allocation into biomolecules by natural phytoplankton assemblages were determined, together with their species composition, in a coastal station of the central Cantabrian Sea (southern Bay of Biscay). Chlorophyll-a concentration ranged from values below 20 mg m⁻² in winter to values above 80 mg m⁻² during spring and during an upwelling event in summer. Low primary production rates (<300 mgC m⁻² d^-1) were measured during winter and during summer stratification periods. The rate of C fixation during summer upwelling conditions exceeded 3500 mgC m⁻² d⁻¹. In terms of photosynthate partitioning, proteins were the dominant fraction, as they typically accounted for >30% of total photo-assimilated C, with polysaccharides and low molecular weight metabolites showing incorporation percentages around 10–30%. Relative C incorporation into lipids was generally <15%. Recurrent patterns of vertical variability in photosynthate partitioning were observed: the relative synthesis of proteins increased toward the bottom of the euphotic zone, whereas the relative C incorporation into polysaccharides and lipids tended to be higher near the surface. When primary production decreased, the synthesis of proteins was maintained more than that of other molecules. Throughout the year, the relative synthesis of proteins was inversely correlated with phytoplankton biomass, production and growth rate. The conservation of protein synthesis under growth-limiting conditions and the enhancement of lipid and polysaccharide synthesis when irradiance is high seem to constitute general patterns of photosynthate partitioning in marine phytoplankton. In our study, these patterns represented metabolic strategies of phytoplankton in response to changing environmental factors, rather than the effect of variations in the species composition of the community.     

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

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