Factors responsible for the differences in satellite-based chlorophyll a concentration between the major global upwelling areas

The aim of this study was to identify the factors responsible for the differences in chlorophyll a concentration (Chl-a) observed between the California, Canary, Humboldt and Benguela upwelling areas. Monthly climatologic values of Chl-a obtained from satellite images, covering the years 1998–2004, revealed that this pigment was higher in the Benguela system than in the other areas. Upwelling intensity, as derived from offshore Ekman transport computations, was higher in the Benguela and Humboldt regions and, for the same upwelling intensity, Chl-a was higher in Benguela than in the other regions. Upwelling intensity appears to be able to drive Chl-a densities through nutrient supply, as nutrients are correlated to offshore Ekman transport. A linear regression model including the fraction of sea surface over the shelf in each 1° × 1° box, nitrate, silicate, turbulence and variability of offshore Ekman transport explained the 84.8% of the variance in Chl-a among the areas. Differences in offshore Ekman transport explained the lower Chl-a observed in Canary and California and the higher Chl-a observed in Benguela and Peru-Humboldt. A narrow continental shelf and low water column stability also contribute to reducing phytoplankton pigment biomass in the Canary and California areas. The higher Chl-a values observed in Benguela compared to Humboldt-Peru are due to a wider extension of the continental shelf in the Benguela region.     

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.     

Zooplankton dynamics in the northern Benguela ecosystem,with special reference to the copepod Calanoides carinatus

Main features of the zooplankton distribution and the ecological characteristics of the dominant species in the northern Benguela during different phases of upwelling are discussed. The composition of the zooplankton between 17 and 27°S was similar each year. Among the 20°30 most abundant species, 3°4 copepods dominated, influencing the distribution of total zooplankton biomass. During quiescent upwelling, zooplankton abundance was low and there were no significant differences in the inshore-offshore distribution of zooplankton biomass, the maximum occurring over the slope. During active upwelling, zooplankton biomass increased significantly, the maximum over the shelf being constituted almost entirely of developmental stages of herbivorous copepods. Over the inner shelf, all stages of the copepod Calanoides carinatus were feeding actively, removing up to 5 per cent per day of the standing stock of phytoplankton. Comparison of daily ration, respiration rate and biochemical composition of C. carinatus revealed active storage of energy inshore. Offshore populations of C. carinatus, found deeper than 200 m, comprised mainly copepodite stage V, which were not feeding and were characterized by decreased mobility and respiration and a high lipid content. It is estimated that the energy stored during active upwelling enables copepods to survive up to six months without any additional source of energy.     

Upwelling and the distribution of chlorophyll a within the Bay of Concepción,Chile

The influence of upwelling on the distribution of chlorophyll a within the Bay of Concepción, Chile is discussed in light of continuous measurements of surface in vivo chlorophyll fluorescence and temperature taken simultaneously along horizontal transects, and hydrographic stations' data. Results suggest significant temporal variability both in the distribution of in vivo fluorescence, temperature and salinity within the Bay and in the characteristics of the exchange between the Bay and the adjacent shelf waters, induced by variable upwelling. Upwelling is produced by the predominant south-westerly winds during the summer. Significant variations in the wind direction occur with periods from two to seven days. During active upwelling, exchange is characterized by a surface outflow through the mouth of the Bay and an inflow at depth. Low chlorophyll fluorescence is confined to the upwelling areas on the eastern shore either within or outside the Bay; high chlorophyll fluorescence is confined to the central and western Bay. Density data suggest a three-layered circulation pattern at the mouth of the Bay during the upwelling relaxation involving an inflow both at the surface and bottom and outflow at mid-depth. Associated with this exchange is an active high chlorophyll transport from the Bay to the adjacent coastal waters at mid-depth and inflow of low chlorophyll water from the adjacent shelf at the surface and near the bottom.     

Summer and winter differences in zooplankton biomass,distribution and size composition in the KwaZulu-Natal Bight,South Africa

Zooplankton biomass and distribution in the KwaZulu-Natal Bight were investigated in relation to environmental parameters during summer (January–February 2010) and winter (July–August 2010). Mean zooplankton biomass was significantly higher in winter (17.1 mg dry weight [DW] m^–3) than in summer (9.5 mg DW m^?3). In summer, total biomass was evenly distributed within the central bight, low off the Thukela River mouth and peaked near Durban. In winter, highest biomass was found offshore between Richards Bay and Cape St Lucia. Zooplankton biomass in each size class was significantly, negatively related to sea surface temperature and integrated nitrate, but positively related to surface chlorophyll a and dissolved oxygen. Zooplankton biomass was significantly related to bottom depth, with greatest total biomass located inshore (<50 m). Distribution across the shelf varied with zooplankton size. Seasonal differences in copepod size composition suggest that a smaller, younger community occupied the cool, chlorophyll-rich waters offshore from the St Lucia upwelling cell in winter, and a larger, older community occurred within the relatively warm and chlorophyll-poor central bight in summer. Nutrient enrichment from quasi-permanent upwelling off Durban and Richards Bay appears to have a greater influence on zooplankton biomass and distribution in the bight than the strongly seasonal nutrient input from the Thukela River.     

Zooplankton distribution and cross-shelf transfer of carbon in an area of complex mesoscale circulation in the northern California Current

We conducted a research cruise in late summer (July–August) 2000 to study the effect of mesoscale circulation features on zooplankton distributions in the coastal upwelling ecosystem of the northern California Current. Our study area was in a region of complex coastline and bottom topography between Newport, Oregon (44.7°N), and Crescent City, California (41.9°N). Winds were generally strong and equatorward for >6 weeks prior to the cruise, resulting in the upwelling of cold, nutrient-rich water along the coast and an alongshore upwelling jet. In the northern part of the study area, the jet followed the bottom topography, creating a broad, retentive area nearshore over a submarine shelf bank (Heceta Bank, 44–44.4°N). In the south, a meander of the jet extended seaward off of Cape Blanco (42.8°N), resulting in the displacement of coastal water and the associated coastal taxa to >100 km off the continental shelf. Zooplankton biomass was high both over the submarine bank and offshore in the meander of the upwelling jet. We used velocities and standing stocks of plankton in the upper 100 m to estimate that 1×10⁶ m³ of water, containing an average zooplankton biomass of ～20 mg carbon m^?3, was transported seaward across the 2000-m isobath in the meandering jet each second. That flux equated to offshore transport of >900 metric tons of carbon each day, and 4–5×10⁴ tons over the 6–8 week lifetime of the circulation feature. Thus, mesoscale circulation can create disparate regions in which zooplankton populations are retained over the shelf and biomass can accumulate or, alternatively, in which high biomass is advected offshore to the oligotrophic deep sea.     

Phytoplankton pigment and absorption characteristics along meridional transects in the Atlantic Ocean

Pigment patterns and associated absorption properties of phytoplankton were investigated in the euphotic zone along two meridional transects in the Atlantic Ocean, between the UK and the Falkland Islands, and between South Africa and the UK. Total chlorophyll a (TChla=MVChla+DVChla+chlorophyllide a) concentrations and the biomarker pigments for diatoms (fucoxanthin), nanoflagellates and cyanobacteria (zeaxanthin) appeared to have similar distribution patterns in the spring and in the autumn in the temperate NE Atlantic and the northern oligotrophic gyre. Divinyl chlorophyll a levels (prochlorophytes) were greater in spring at the deep chlorophyll maximum in the oligotrophic gyre, however. Marked seasonal differences were observed in the NW African upwelling region. TChla concentrations were twice as high in the upper mixed layer in the spring, with the community dominated by diatoms and prymnesiophytes (19′-hexanoyloxyfucoxanthin). A layered structure was prevalent in the autumn where cyanobacteria, diatoms and prymnesiophytes were located in the upper water column and diatoms and mixed nanoflagellates at the sub-surface maximum. In the South Atlantic, the Benguela upwelling ecosystem and the Brazil-Falklands Current Confluence Zone (BFCCZ) were the most productive regions with the TChla levels being twice as high in the Benguela. Diatoms dominated the Benguela system, while nanoflagellates were the most ubiquitous group in the BFCCZ. Pigment concentrations were greater along the eastern boundary of the southern oligotrophic gyre and distributed at shallower depths. Deep chlorophyll maxima were a feature of the western boundary oligotrophic waters, and cyanobacteria tended to dominate the upper water column along both transects with a mixed group of nanoflagellates at the chlorophyll maximum.Absorption coefficients were estimated from spectra reconstructed from pigment data. Although absorption was greater in the productive areas, the TChla-specific coefficients were higher in oligotrophic regions. In communities that were dominated by diatoms or nanoflagellates, pigment absorption was generally uniform with depth and attenuating irradiance, with TChla being the major absorbing pigment at 440 nm and photosynthetic carotenoids (PSC) at 490 nm. Absorption by chlorophyll c and photoprotective carotenoids (PPC) was much lower. Populations where cyanobacteria were prevalent were characterized by high PPC absorption, particularly at 490 nm, throughout most of the euphotic zone. The data suggested that the effect of pigments on the variability of phytoplankton absorption was due primarily to the variations in absorption by PPC.     

Environmental influence on phytoplankton communities in the northern Benguela ecosystem

An investigation of surface phytoplankton communities was undertaken on the shelf of the northern Benguela upwelling ecosystem during austral autumn (May) and spring (September), along latitudinal transects at 20° S and 23° S, from 2 to 70 nautical miles offshore, as well as on a zigzag grid located between these transects. Microscopic identification of the phytoplankton and CHEMTAX analysis of pigment biomarkers were used to characterise the community composition. During May 2014, warmer, more-saline water with a shallower upper mixed layer corresponding to periods of less-intense offshore Ekman transport was encountered on the shelf. Satellite imagery indicated high phytoplankton biomass extending for a considerable distance from the coast, and CHEMTAX indicated diatoms as dominant at most of the stations (52–92%), although dinoflagellates were dominant at some inshore localities (57–74%). Species of Chaetoceros, Bacteriastrum and Cylindrotheca were the most abundant, with abundance of the Pseudo-nitzschia ‘seriata-group’ being particularly high at a number of stations. In September 2014, more-intense wind-forcing resulted in a deeper upper mixed layer and stronger upwelling of colder, less-saline water. Elevated phytoplankton biomass was confined close to the coast, where diatoms accounted for most of the population (54–87%), whereas small flagellates, such as prasinophytes, haptophytes and cryptophytes, as well as the cyanobacterium Synechococcus, dominated the communities (58–90%) farther from the coast. It is hypothesised that stronger upwelling and deeper vertical mixing in September of that year were not conducive for widespread diatom growth, and that small flagellates populated the water column by being entrained from offshore onto the shelf in the upwelled water that moved in towards the coast.     

Zooplankton and the oceanography of the eastern tropical Pacific: A review

We review the spatial and temporal patterns of zooplankton in the eastern tropical Pacific Ocean and relationships with oceanographic factors that affect zooplankton distribution, abundance and trophic relationships. Large-scale spatial patterns of some zooplankton groups show broad coincidence with surface water masses, circulation, and upwelling regions, in agreement with an ecological and dynamic partitioning of the pelagic ecosystem. The papers reviewed and a new compilation of zooplankton volume data at large-scale show that abundance patterns of zooplankton biomass have their highest values in the upwelling regions, including the Gulf of Tehuantepec, the Costa Rica Dome, the equatorial cold tongue, and the coast of Peru.Some of the first studies of zooplankton vertical distribution were done in this region, and a general review of the topic is presented. The possible physiological implications of vertical migration in zooplankton and the main hypotheses are described, with remarks on the importance of the oxygen minimum zone (OMZ) as a barrier to both the vertical distribution and migration of zooplankton in the region. Recent results, using multiple-net gear, show that vertical distribution is more complex than previously thought. There are some well-adapted species that do live and migrate within the OMZ.Temporal patterns are reviewed and summarized with historical data. Seasonal variations in zooplankton biomass follow productivity cycles in upwelling areas. No zooplankton time series exist to resolve ENSO effects in oceanic regions, but some El Niño events have had effects in the Peru Current ecosystem. Multidecadal periods of up to 50 years show a shift from a warm sardine regime with a low zooplankton biomass to a cool anchovy regime in the eastern Pacific with higher zooplankton biomasses. However, zooplankton volume off Peru has remained at low values since the 1972 El Niño, a trend opposite to that of anchoveta biomass since 1984.Studies of trophic relations emphasize the difference in the productivity cycle in the eastern tropical Pacific compared to temperate or polar ecosystems, with no particular peaks in the stocks of either zooplankton or phytoplankton. Productivity is more dependent on local events like coastal upwelling or water circulation, especially in the equatorial countercurrent and around the equatorial cool-tongue. Micrograzers are very important in the tropics as are predatory mesozooplankton. Up to 70% of the daily primary productivity is consumed by microzooplankton, which thus regulates the phytoplankton stocks. Micrograzers are an important link between primary producers, including bacteria, and mesozooplankton, constituting up to 80% of mesozooplankton food. Oceanography affects zooplankton trophic relationships through spatial–temporal effects on primary productivity and on the distributions of metabolic factors, food organisms, and predators. This paper is part of a comprehensive review of the oceanography of the eastern tropical Pacific.     

Spatial and seasonal variation in chlorophyll distribution in the upper 30 m of the photic zone in the southern Benguela/Agulhas ecosystem

Depth-integrated chlorophyll a in the upper 30 m is used as an index of phytoplankton biomass. Mean concentrations of chlorophyll a (1971–1989) were calculated for half-degree rectangles of latitude and longitude within the 500 m isobath off the South African coast. These data were used to estimate median and mean concentrations and coefficients of variance for different seasons and geographical strata (i.e. inshore and offshore regions of the continental shelf along the West, Cape and South coasts). Offshore, longshore and seasonal differences in the distribution of phytoplankton biomass in the Benguela/Agulhas system were tested for statistical significance.     

Analysis of temporal and spatial variability of phytoplankton by physical-biological models

I reviewed my research on analysis of temporal and spatial variability of phytoplankton by physical-biological models. This paper was prepared for a lecture of the member awarded the Okada Prize for 1991 from the Oceanographical Society of Japan.Temporal change of phytoplankton in a local upwelling was studied by simulated upwelling experiments conducted with natural phytoplankton communities under natural surface light conditions. Results of the culture experiments was explained by a simple model. This model allows to predict the chlorophyll and nutrient concentration changes in a given upwelled water mass.Above model was verified by a local upwelling observed off Izu, Japan, on May, 1982. Phytoplankton growth and nutrient decrease in surface water of the local upwelling were observed within two days followed by decrease of phytoplankton concentration under depleted nutrient environment. The phytoplankton growth and nutrient decrease could explained by the model with phytoplankton removal rate of about half of the growth rate. Centric diatom was the dominant phytoplankton group and pennate diatom showed less abundance in the upwelled water. Pennate diatom showed fast growth rate when nutrient was abundant and fast decreasing rate after nutrient depleted. On the other hand, flagellate and monads showed relatively slow change of biomass under the change of nutrient concentrations. Furthermore, resting spore formation of centric diatom,Leptocylindrus danicus, was observed in a response to nutrient depletion.Temporal and spatial variability of phytoplankton in the southeastern U.S. continental shelf ecosystem was studied by physical-biological models. First, differences of the biological responses to frontal eddy upwelling during spring and to intrusion during summer was considered by Lagrangian particle tracing experiments with optimally-interpolated flow fields. In spring, particles showed residence time of a few days; however, particles in summer intrusion stayed on the shelf nearly 30 days. It was concluded that difference of particle residence time of upwelled water make the difference of plankton communities. Similar flow fields and particle tracing experiments were used to trace the features in chlorophyll distributions during spring of 1980 derived by Coastal Zone Color Scanner (CZCS). Phytoplankton patchness were created and deformed by frontal eddy events. Eularian physical-biological model was constructed to understand the CZCS-chlorophyll distributions. Statistical comparisons with series of numerical experiments indicate that horizontal advection is an important process for the chlorophyll distributions and that upwelling and associated phytoplankton growth are responsible for the across-shelf gradients and maintenance of concentrations. Furthermore, the CZCS data were assimilated to the model to improve the phytoplankton concentrations, and phytoplankton carbon flux across shelf was estimated. Processes causing the time changes of chlorophyll concentrations were estimated with the model and satellite data further indicated that the both physical and biological forcing is important for the time chages. Several other studies conducted presently were mentioned.     

北冰洋浮游生物空间分布及其季节变化的模拟

低营养级浮游生物生态动力过程对环境变化的响应非常敏感。随着全球气候变化加剧，北冰洋正在经历快速的环境变化。厘清北冰洋低营养级浮游生物季节分布与变化特征是探究北冰洋生态系统对环境快速变化响应的前提，也是评估北极海区固碳能力的重要依据。基于此，本文构建了海洋–海冰–生物地球化学循环模型，并对北冰洋叶绿素浓度以及浮游生物结构的时空变化特征进行了模拟，结果表明:(1)北冰洋表层叶绿素浓度的峰值主要出现在5月，且太平洋一侧叶绿素浓度高于大西洋一侧；随着海水层化，表层受营养盐限制的海区呈现次表层叶绿素浓度最大值现象，且由陆架向海盆，次表层叶绿素浓度最大值层逐渐加深；9月，叶绿素浓度高值重回水体上层，太平洋一侧海区表层叶绿素浓度呈现较为明显的次峰值。(2)由于太平洋和大西洋入流营养盐浓度及结构的不同，北冰洋表层浮游生物群落结构存在明显空间差异。太平洋一侧，硅藻和中型浮游动物占优，硅藻在5月和9月出现生物量峰值，微型浮游植物在3月、5月和6月维持相对较高生物量；而大西洋一侧，在早春－春末夏初－夏秋经历了微型浮游植物－硅藻－微型浮游植物的演替，总体而言，微型浮游植物和微型浮游动物占优。此外，两侧海区浮游动物浓度峰值相较浮游植物滞后约半月。     

Seasonal Variations of Nutrients, Seston and Phytoplankton, and Upwelling Intensity off La Coruña (NW Spain)

This study describes the main seasonal stages in oceanographic conditions and phytoplankton off La Coruña (Galicia, NW Spain), during 1991 and 1992, based mainly on monthly cruises near the coast. Upwelling conditions were studied using an upwelling index calculated from local winds. The Galician coast is affected by a long upwelling season for most of the year. The upwelling pulses interact with the thermal stratification-mixing cycle of surface waters, primarily affecting the dynamics of phytoplankton. In addition, the presence of water masses of different salinity in the subsurface layers changes the stratification of the water column. The less-saline North Atlantic Central Water (NACW) was normally associated with upwelling events during summer. However, on several occasions during the study, the presence of Eastern North Atlantic Water (ENAW) of subtropical origin was observed with salinities up to 36·22 and temperatures between 13 and 14 °C.Observations were grouped into five main stages related to the degree of surface stratification and characteristics of phytoplankton communities. These stages were recognized in both annual cycles, and were termed: winter mixing, spring and autumn blooms, summer upwelling, thermal stratification and special events (red tides and downwelling). A homogeneous water column was the main characteristic of the winter stage, with high nutrient concentrations and low phytoplankton biomass. Eastern North Atlantic Water appeared at the end of this stage, which lasted from November to February. The spring and autumn blooms occurred along with weak thermohaline gradients at the surface, producing high phytoplankton concentrations. Favourable upwelling conditions and the presence of ENAW in a subsurface layer were the factors that most likely induced earlier blooms, while thermal gradients developed at the surface could have been more important for later blooms. Upwelling events during summer were related to a reduction in the depth of the surface mixed layer as the pycnocline moved upwards, and can produce significant phytoplankton accumulations. These summer blooms interrupted the thermal stratification stage, characterized by low nutrient and phytoplankton concentrations at the surface. The dominant phytoplankton in the study was composed mainly of diatoms, especially during blooms. However, a proliferation of red-tide dinoflagellates was observed along with weak upwelling conditions in late summer. Also in late summer, strong downwelling conditions caused the accumulation of warmer shelf waters inshore, inducing the sinking of particulate matter produced at the surface.     

Long-term changes of dissolved oxygen,hypoxia, and the responses of the ecosystems in the East China Sea from 1975 to 1995

Observation data obtained in the 32°N transect (transect E) in 1975–1995 were used to analyze the long-term changes in dissolved oxygen (DO) concentration and near-bottom hypoxic water in the East China Sea (ECS). A declining trend in annual average DO concentration and the degree of DO saturation was observed. Consequently, the apparent oxygen utilization in the western waters of transect E was on the rise. There was a seasonal hypoxic phenomenon in near-bottom water in the western water of transect E. The width of hypoxic water formed in summer gradually extended eastward along the continental shelf (transect E) at the rate of 3.12 km year⁻¹. Three potential reasons might have caused the formation and maintenance of near-bottom hypoxic water. First, the special hydrological topography and hypoxic deep water of the Taiwan Warm Current provided a backdrop for the hypoxic zone. Second, in summer, the strength of water column stratification restricts water exchange. Third is the occurrence and decay of the phytoplankton bloom. In surface water, nutrient concentrations increased gradually, and chlorophyll (Chl a), primary production, and phytoplankton biomass in summer increased. On the other hand, the community structure of phytoplankton, zooplankton, and zoobenthos became simple. Blooming phytoplankton consumed plenty of nutrients in the surface, but the upwelling of nutritious bottom water was suppressed by the strong thermocline. As a result, sinking of phytoplankton was enhanced because of nutrient deficiency. In recent years, a serious lack of zoobenthos in the study area corresponded to a higher degree of hypoxia. This phenomenon would have a major effect on the evolution of ecological dynamic systems in the ECS.     

On the size of the Peru upwelling ecosystem

Previously published estimates of the area of the Peru upwelling ecosystem vary by more than an order of magnitude. In an effort to improve this situation, we used a 24-month sequence of SeaWiFS satellite images of chlorophyll in the surface water off Peru from 5°S to 18.5°S during September 1997–August 1999 to estimate the size of the nutrient enhanced productive habitat associated with the upwelling. The first 12-month period was marked by El Niño conditions, the second by strong upwelling. Using a chlorophyll threshold of >1.0 mg m⁻³ to define the limit of the productive habitat resulted in maximum area estimates of 120×10³ km² during September 1997–August 1998, and 220×10³ km² during September 1998–August 1999. The latter result is consistent with an area estimate we calculated using total fishery landings and a regression relating fishery yields per unit area to annual primary production per unit area. Although year-to-year variation in the annual mean size of the upwelling ecosystem must be significant, even discounting El Niño events, our analysis has shown that at least five of the extreme earlier values are not good estimates of the size of the productive habitat. We may now be close to knowing the average size of the ecosystem to within a factor of about two.     

Inshore-offshore and vertical patterns of phytoplankton biomass and community composition in Santa Monica Bay, CA (USA)

Spatial gradients in biomass and community composition have important consequences for ecosystem structure and function. In this study, small-scale inshore-offshore (1-10 km) and vertical (1-50 m) patterns of microphytoplankton biomass and community composition are described, and the environmental controls of microphytoplankton biomass are evaluated in a coastal ecosystem of the Southern California Bight (SCB). During a two-year period, persistent inshore-offshore gradients in phytoplankton biomass and occasional inshore-offshore gradients in community composition, coincident with regional precipitation, were found, although the strength of the gradients varied between sampling periods. The chlorophyll a maximum was generally present between 15 and 45 m, the cell abundance maximum occurred in surface waters, and there was little evidence of vertical gradients in community composition. Variability in chlorophyll a concentrations was linked to variability in environmental parameters only after some rain and upwelling events. This study demonstrates that inshore-offshore patterns in phytoplankton biomass previously documented at large spatial scales (100-700 km) in the SCB can also persist at smaller scales (1-10 km), although the mechanisms for the gradients are likely to be different at the different spatial scales. The results provide a baseline data set that can be used to focus monitoring and management efforts in the SCB. In particular, this work shows that a limited number of sampling stations are sufficient for phytoplankton monitoring in Santa Monica Bay.     

Monsoonal influence on the distribution of phytoplankton pigments in the Arabian Sea

Variations in the distribution of chemotaxonomic pigments were monitored in the Arabian Sea and the Gulf of Oman at the end of the SW monsoon in September 1994 and during the inter-monsoon period in November/December 1994 to determine the seasonal changes in phytoplankton composition. The Gulf of Oman was characterized by sub-surface chlorophyll maxima at 20-40 m during both seasons, and low levels of divinyl chlorophyll a indicated that prochlorophytes did not contribute significantly to the total chlorophyll a. Prymnesiophytes (19′-hexanoyloxyfucoxanthin), diatoms (fucoxanthin) and chlorophyll b containing organisms accounted for most of the phytoplankton biomass in September, while prymnesiophytes dominated in November/December. In the Arabian Sea in September, high total chlorophyll a concentrations up to 1742 ng l^-1 were measured in the coastal upwelling region and a progressive decline was monitored along the 1670 km offshore transect to oligotrophic waters at 8°N. Divinyl chlorophyll a was not detected along this transect except at the two most southerly stations where prochlorophytes were estimated to contribute 25–30% to the total chlorophyll a. Inshore, the dominance of fucoxanthin and/or hexanoyloxyfucoxanthin indicated that diatoms and prymnesiophytes generally dominated the patchy phytoplankton community, with zeaxanthin-containing Synechococcus also being important, especially in surface waters. At the southern oligotrophic localities, Synechococcus and prochlorophytes dominated the upper 40 m and prymnesiophytes were the most prominent at the deep chlorophyll maximum. During the inter-monsoon season, total chlorophyll a concentrations were generally half those measured in September and highest levels were found on the shelf (1170 ng l^-1). Divinyl chlorophyll a was detected at all stations along the Arabian Sea transect, and we estimated that prochlorophytes contributed between 3 and 28% to the total chlorophyll a, while at the two oligotrophic stations this proportion increased to 51–52%. While procaryotes were more important in November/December than September, eucaryotes still accounted for >50% of the total chlorophyll a. Pigment/total chlorophyll a ratios indicated that 19′-hexanoyloxyfucoxanthin-containing prymnesiophytes were the dominant group, although procaryotes accounted for 65% at the two southerly oligotrophic stations.     

St Helena Bay (southern Benguela) then and now: muted climate signals,large human impact

The development of suitable reference states for ecosystem-based management requires documentation of changes in structure and functioning of marine ecosystems, including assessment of the relative importance of bottom-up and top-down processes as drivers of change. We used monitoring data available from St Helena Bay, the most productive bay and an important nursery area situated on the west coast of South Africa, during 1950–2010 to reveal changes in the abiotic and biotic components. St Helena Bay in the 1950s showed similarities to 2000–2010 in terms of wind patterns, hydrology and phytoplankton. Upwelling, oxygen and nutrient concentrations in subthermocline water displayed pronounced decadal-scale variability. Primary production in St Helena Bay is variable, but consistently higher than that on the adjacent Namaqua shelf. Zooplankton size composition and biomass in August have changed markedly since the 1950s. During 2001–2010, mesozooplankton biomass in autumn was considerably lower than in summer, probably due to predation by small pelagic fish. Pelagic fish catch patterns and distributions have altered dramatically. Conservation measures, implemented to reverse past negative human impact, have benefitted marine mammals, the abundance of which has increased in the area, but additional conservation measures are necessary to reverse the decline in African penguins Spheniscus demersus. St Helena Bay shows a muted response to long-term change in the southern Benguela, with marked decadal variability but no clear long-term trend in oceanography and biogeochemistry. Changes in ecosystem boundary conditions and fishing pressure cannot be ignored as important drivers of change in the southern Benguela since the 1950s.     

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

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

The Western Boundary of the Equatorial Pacific Upwelling: Some Consequences of Climatic Variability on Hydrological and Planktonic Properties

The longitude of the western limit of the equatorial Pacific upwelling is a key parameter for studies of carbon budget and pelagic fisheries variability. Although it is well defined at the surface on the equator by a salinity front and a sharp variation of the partial pressure of CO₂, data from two equatorial cruises make it clear that this hydrological limit does not necessarily coincide with the boundary of the nitrate and chlorophyll enriched area. In January-February 1991 during a non-El Niño period, when trade winds and the South Equatorial current (SEC) were favorable to upwelling, the two limits were at the same longitude. Conversely, in September-October 1994 during El Niño conditions, when the equatorial upwelling had stopped, the nitrate and chlorophyll enriched zone was found a few degrees of longitude east of the hydrological boundary (5.5° at the surface and 2.5° for the 50 m upper layer), whereas no such offset was observed for zooplankton biomass. A simple model, based on the HNLC (High Nutrient - Low Chlorophyll) ecosystem functioning, was initialized with nitrate uptake measurements and estimates of upwelling break duration. The model results support the hypothesis that zonal separation of the limits arises from biological processes (i.e. nitrate uptake and phytoplankton grazing) achieved during that upwelling break.