Nanoplanktonic assemblages in the upwelling area off Concepción (∼36°S), central Chile: Abundance, biomass, and grazing potential during the annual cycle

The structure and functioning of nanoplanktonic assemblages in coastal upwelling areas have usually been overlooked in explorations of the productivity of these areas. As part of a multidisciplinary, time-series station in the coastal area off Concepción, seasonal variations (upwelling and non-upwelling) in the abundance and biomass of these assemblages were investigated. Hydrographic measurements and biological samples were taken monthly over a 2-year period (18 August 2004-28 July 2006). Nanoflagellates dominated the total integrated abundance (3-317 × 10⁹ cells m⁻²; 0-80 m). Diatoms and dinoflagellates usually contributed to a lesser degree (<20%) but sporadically made important contributions to the total integrated nanoplankton biomass (0.02-10.6 g C m⁻²). Most of the nanoplankton was concentrated in surface waters (<30 m) during all the samplings and no seasonal differences in abundance or biomass were found in this layer, although the mean values and dispersions around them were highest during the upwelling period along with maximum integrated (0-80 m) chlorophyll-a values, as total or in the <20 μm fraction. Changes in nanoplankton abundance were significantly but weakly (r < 0.4) correlated with changes in the hydrographic variables; the highest correlation values were positive for temperature and oxygen, factors that varied with depth and date. The potential grazing rates of heterotrophic nano-predators (flagellates and dinoflagellates) on prokaryotic prey, estimated with a generic model, ranged from 3 to 242 bacterioplankton predator⁻¹ h⁻¹ and from 0.1 to 14 cyanobacteria predator⁻¹ h⁻¹. Our results imply a small impact of seasonal hydrographic variability on the abundance and biomass of nanoplanktonic assemblages and suggest that grazing by nanoheterotrophs might control the prokaryotic picoplankton populations in the upwelling area off Concepción.     

The contribution of nano- and micro-planktonic assemblages in the surface layer (0-30 m) under different hydrographic conditions in the upwelling area off Concepción, central Chile

In the highly productive region off central Chile, the structure and temporal and spatial variability of planktonic assemblages, and the factors that determine changes in this structure are poorly understood. In the region, wind-driven upwelling, heating by solar radiation and freshwater inputs are highly seasonal processes, which, together with higher frequency events, can promote changes in the planktonic communities, especially in the upper layer. This study focuses on the structure of nano- through to micro-planktonic assemblages (2-200 μm) of unicellular organisms (protists) in surface waters (0-30 m) during different hydrographic conditions. Samples were taken from a fixed shelf station off Concepción (COPAS time series Station 18) on eight occasions between September 2003 and August 2004. The nano-plankton flagellate-dominated fraction was numerically important during the whole period. Maxima in flagellate abundance and biomass occurred during the upwelling period (November-April samplings) but these maxima appear to be unrelated to the degree of water column stratification. The micro-plankton diatom-dominated fraction was usually the largest component in terms of biomass during the study period and the diatoms made important numerical contributions during the upwelling period, with maxima in abundance and biomass when water column stability was lowest. The dominant genera and morphotypes in each functional group were found throughout the study period, with maxima in abundance and biomass co-occurring under similar environmental upwelling conditions. The mean macro-nutrient concentrations (nitrate and silicate) were relatively high in the top 30 m during both upwelling and non-upwelling periods, and did not explain the maxima in plankton or functional group replacements. The persistence of the dominant taxa in the planktonic assemblages suggests a high degree of flexibility, though probably not at the specific level, to withstand the highly variable environmental conditions in this upwelling area.     

The Influence of Winds and Tides in the Formation of Circulation Layers in a Bay, a Case Study: Concepción Bay, Chile

Horizontal water velocities were measured using three current meters moored at the levels of 5, 18, and 30 m depth in the oriental side of Concepción Bay (36°40′S; 73°02′W), an almost rectangular, shallow bay off central Chile, connected with the open sea through two mouths.The large momentum transfer from the wind to the water causes variations of the internal volume of the bay generating circulation layers having almost anti-parallel directions. Northerly winds carry superficial water into the bay, causing a compensating flow leaving the bay near its bottom. Winds from the SW, on the other hand, cause an outgoing circulation layer in the surface and a circulation layer entering into the bay near the bottom. During wind calm periods, longer than 12 h, the two layers disappear, leaving a current system comprising only one circulation layer. The response time necessary for the generation or dissipation of these layers, which fluctuates between 1 and 7 h, depends on the initial conditions of the bay and on the intensity, direction and persistence of the wind.     

Productivity cycles in the coastal upwelling area off Concepción: The importance of diatoms and bacterioplankton in the organic carbon flux

Recurrent coastal upwelling is recognized as one of the main factors promoting the exceptionally high productivity of the Humboldt Current System. Herein, we study time series data of gross primary production (2003-2006) and its fluctuation in relation to seasonal changes in the light and nutrient field of the Concepción upwelling ecosystem. Concurrent measurements of gross primary production, community respiration, bacterial secondary production, and sedimentation rates allowed a characterization of the main carbon fluxes and pathways in the study area. The integrated values of gross primary production were higher during the upwelling period (>1 g C m⁻² d⁻¹; October-April; that is, early spring to early austral fall). Seasonal changes in the system were also reflected in community respiration, organic matter sedimentation, and bacterial production rates, which varied along with the gross primary production. The significant correlation between gross primary production and community respiration (Spearman, r = 0.7; p < 0.05; n = 18) reflected an important degree of coupling between organic matter formation and its usage by the microplanktonic community during periods when gross primary production/community respiration were highly similar. Higher gross primary production values (>6 g C m⁻² d⁻¹) were consistently associated with maximum biomass levels of Skeletonema costatum and Thalassiosira subtilis. We observed a positive correlation between gross primary production and the sedimentation of intact diatom cells (Spearman, r = 0.5, p < 0.05, n = 17). Our data suggest that, in the Concepción upwelling ecosystem, bacteria utilize an important fraction of the gross primary production. If our interpretations are correct, they leave unanswered the question of how the system supports the extremely high fish biomass levels, therein pointing out the system’s limited capacity to buffer the evasion of CO₂ following upwelling.     

Temporal variability of Pb fluxes and bioturbation in shelf sediments beneath the high primary production area off Concepción, central-southern Chile (36°S)

We estimated monthly fluxes of ²¹⁰Pb in shelf sediments beneath a high productivity area off central-southern Chile (36°S) during 1 year (September 2002-August 2003). Sediment cores were obtained using a multiple corer and were analyzed mainly for ²¹⁰Pb, total pigments, and macrofauna abundance. The ²¹⁰Pb inventories and fluxes were estimated for surface sediments (0-5 cm) and bioturbation coefficients were inferred using chlorophyll-a (reactive) profiles. In general, ²¹⁰Pb content was inversely correlated with phytodetritus fluxes. High photosynthetic pigment contents in surface sediments were consistently associated with lower ²¹⁰Pb contents. Macrofaunal activity responded to oxygen and organic matter supplies at the sediment surface, generally concentrated in the first centimeters, but particularly so during months of high organic matter fluxes and deficient bottom water oxygen conditions. At this study site, several processes involved in the ²¹⁰Pb surface distribution make it difficult to accurately estimate ages at the surface. We postulate that the organic fluxes promote changes in the faunal activity, which, in combination with sediment resuspension and water circulation over the shelf, produce seasonal variations in the ²¹⁰Pb inventories.     

Seasonal and inter-annual variation of mesozooplankton in the coastal upwelling zone off central-southern Chile

Zooplankton sampling at Station 18 off Concepción (36°30′S and 73°07′W), on an average frequency of 30 days (August 2002 to December 2005), allowed the assessment of seasonal and inter-annual variation in zooplankton biomass, its C and N content, and the community structure in relation to upwelling variability. Copepods contributed 79% of the total zooplankton community and were mostly represented by Paracalanus parvus, Oithona similis, Oithona nana, Calanus chilensis, and Rhincalanus nasutus. Other copepod species, euphausiids (mainly Euphausia mucronata), gelatinous zooplankton, and crustacean larvae comprised the rest of the community. Changes in the depth of the upper boundary of the oxygen minimum zone indicated the strongly seasonal upwelling pattern. The bulk of zooplankton biomass and total copepod abundance were both strongly and positively associated with a shallow (<20 m) oxygen minimum zone; these values increased in spring/summer, when upwelling prevailed. Gelatinous zooplankton showed positive abundance anomalies in the spring and winter, whereas euphausiids had no seasonal pattern and a positive anomaly in the fall. The C content and the C/N ratio of zooplankton biomass significantly increased during the spring when chlorophyll-a was high (>5 mg m⁻³). No major changes in zooplankton biomass and species were found from one year to the next. We concluded that upwelling is the key process modulating variability in zooplankton biomass and its community structure in this zone. The spring/summer increase in zooplankton may be largely the result of the aggregation of dominant copepods within the upwelling region; these may reproduce throughout the year, increasing their C content and C/N ratios given high diatom concentrations.     

Similar potential ATP-P production and enzymatic activities in the microplankton community off Concepción (Chile) under oxic and suboxic conditions

The effects of the oxygen minimum zone on the metabolism of the heterotrophic microplankton community (0.22-100 μm) in the Humboldt Current System, as well as the factors controlling its biomass production, remain unknown. Here we compare the effect of four sources of dissolved organic carbon (glucose, oxaloacetate, glycine, leucine) on microbial biomass production (such as ATP-P) and the potential enzymatic activities involved in catabolic pathways under oxic and suboxic conditions. Our results show significant differences (p < 0.05) in the ATP-P production when induced by the different substrates that are used as dissolved organic carbon herein. The induction of ATP-P production is enhanced from glucose < oxaloacetate < glycine < leucine. Nevertheless, for individual substrates, no significant differences were found between incubation under oxic and suboxic conditions except in the case of leucine. For this amino acid, the induction of ATP-P synthesis was higher under suboxic than oxic conditions. The data sets of all the substrates used showed greater potential ATP-P production under suboxic than oxic conditions. The results of the potential enzymatic activities suggest that malate dehydrogenase has the highest signal of NADH oxidization activity in the microbial assemblage. Furthermore, for all experiments, the malate dehydrogenase activity data set had a significant relationship with ATP-P production. These findings suggest that the microbial community inhabiting the oxygen minimum zone has the same or greater potential growth than the community inhabiting more oxygenated strata of the water column and that malate dehydrogenase is the activity that best represents the metabolic potential of the community.     

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.     

Biogeochemistry of surface sediments off Concepción (∼36°S), Chile: El Niño vs. non-El Niño conditions

We compared the signals of several water column properties (upwelling intensity, sea level anomaly, temperature, nutrients, dissolved oxygen, chlorophyll-a, and surface sediments) of the continental shelf off Concepción (36°S) during the 1997-1998 El Niño with those of a normal year (2002-2003). We found that the primary hydrographic effect of El Niño 1997-1998 was a reduction in the input of nutrient-rich, oxygen-poor Equatorial Subsurface Water over the shelf. This affected the biology of the water column, as evidenced by the reduced phytoplankton biomass. Surface sediment properties (biogenic opal, organic carbon, bulk δ¹⁵N) observed during El Niño 1997-1998 reflected a reduced export production and the sediments failed to show the water column seasonality that occurs under normal conditions. In addition, weakened denitrification and/or upper water column fertilization could be inferred from the sedimentary δ¹⁵N. Although diminished, export production was preserved in the surface sediments, revealing less degraded organic matter in the upwelling period of the El Niño year than in the normal year. We suggest that the fresher organic material on the seafloor was probably associated with a severe reduction in the polychaete Parapronospio pinnata, which is considered to be the most important metazoan remineralizer of organic carbon at the sediment-water interface in the study area.     

Fine-scale vertical distribution of coastal and offshore copepods in the Golfo de Arauco, central Chile, during the upwelling season

In order to understand the mechanism by which zooplankters from different origins co-occur during the upwelling season within Golfo de Arauco, one of the most productive areas in central Chile, we assessed short term variations in the vertical distribution of the most abundant copepod species. Fine-scale, day and night vertical zooplankton sampling was done with a pump over 12 days in summer. The water column in the gulf consisted of three layers: Equatorial Subsurface Water of low dissolved oxygen content in the deeper part of the water column, strong temperature and oxygen gradients at mid-depth (15-25 m), and a layer of warmer, more oxygenated, less saline water at the surface. Copepods within the gulf originated from offshore, from the continental shelf, and from the coastal area. Most taxa showed distinctive vertical distributions. Three copepod groups were identified by their mean weighted depths of residence. One group included shallow residents found above the thermocline/oxycline (Acartia tonsa, Centropages brachiatus, Corycaeus sp., Paracalanus parvus, Oncaea sp.). A second group was comprised by species distributed at or below the thermocline/oxycline (Oithona sp., Oncaea conifera, Lucicutia sp., Metridia sp., Heterorhabdus papilliger). The third group was composed of vertical migrators that crossed the thermocline/oxycline (Calanus chilensis, Calanoides patagoniensis, Aetideus armatus, Pleuromamma piseki). In spite of their different vertical distribution ranges, the most abundant and frequent copepod species (P. parvus, C. chilensis, C. patagoniensis, C. brachiatus) share a common capacity to withstand wide ranges of oxygen concentration and temperature. This characteristic, along with the capacity to vary their life strategies under different environmental conditions, seems to facilitate the maintenance of large numbers of copepods in coastal waters along the Humboldt Current.     

Spatial and temporal variability of microplankton and detritus, and their export to the shelf sediments in the upwelling area off Concepción, Chile (∼36°S), during 2002-2005

Seasonal variations in diversity and biomass of diatoms, tintinnids, and dinoflagellates and the contribution of microplankton and faecal material to the vertical flux of particulates were investigated at one time series station T (station 18) between 2002 and 2005 and at a grid of stations during November 2004 in the coastal and oceanic area off Concepción (36°S), Chile. The variations were analysed in relation to water column temperature, dissolved oxygen, nutrient concentration, offshore Ekman transport, and chlorophyll-a concentration. Abundance was estimated as cell numbers per litre and biomass in terms of biovolume and carbon units.A sharp decrease with depth was observed in the abundance of both phytoplankton and microzooplankton during the whole annual cycle; over 70% of their abundance was concentrated in the upper 10 m of the water column. Also, a clear seasonality in microplankton distribution was observed at station T, with maxima for diatoms, tintinnids, and dinoflagellates every summer (centred on January) from 2002 to 2005.On the grid of stations, the maximum integrated (0-50 m) micro-phytoplankton abundances (>1 × 10⁹ cells m⁻²) occurred at the coastal stations, an area directly influenced by upwelling. A similar spatial distribution was observed for the integrated (0-200 m) faecal carbon (with values up to 632 mg C m⁻²). Tintinnids were distributed in all the first 300 miles from the coast and dinoflagellates were more abundant in oceanic waters.At station T, the average POC export production (below 50 m depth) was 16.6% (SD = 17%; range 2-67%; n = 16). The biological-mediated fluxes of carbon between the upper productive layer and the sediments of the continental shelf off Concepción depend upon key groups of phytoplankton (Thalassiosira spp., Chaetoceros spp.) and zooplankton (euphausiids) through the export of either cells or faecal material, respectively.     

Strong CO2 outgassing from high nutrient low chlorophyll coastal waters off central Chile (30°S): The role of dissolved iron

Carbonate system parameters (pH and alkalinity) were used to estimate the coastal water CO₂ fluxes off central Chile (30°S) during September 2007. Coastal waters rich in nitrate and silicate were strongly CO₂ supersaturated and normally poor in chlorophyll a. MODIS satellite chlorophyll a data suggest that phytoplankton biomass remained particularly low during September 2007 although coastal waters were highly fertilized with nitrate and silicate. The phytoplankton gross primary productivity in macronutrient-rich waters was very low with the exception of shallow waters (e.g. within or near bays). Several iron-enrichment bottle experiments show that fCO₂ rapidly decreases during iron-enrichment treatments compared to controls. This suggests that iron limitation of phytoplankton growth (mainly diatoms) plays a role in maintaining high-CO₂ outgassing by preventing rapid interception of upwelled CO₂.     

Abundance and distribution of pelagic piscivorous fishes in the Columbia River plume during spring/early summer 1998-2003: Relationship to oceanographic conditions, forage fishes, and juvenile salmonids

From 1998 to 2003, we observed large fluctuations in the abundance and distribution of four pelagic predatory (piscivorous) fishes off northern Oregon and southern Washington, USA. Fluctuations in predatory fish species composition and abundance were strongly linked to the date of the spring transition and to ocean temperatures. Predatory fishes, forage fishes, and juvenile salmonids had distinct spatial distributions, with predators distributed primarily offshore and forage fish and salmonids onshore, but this varied depending on ocean conditions. We suggest that predatory and forage fish distributions respond to ocean temperatures, predator/prey interactions, and possibly turbidity. A shift in ocean conditions in 1999 decreased overall predator fish abundance in the Columbia River plume, particularly for Pacific hake. Marine survival of juvenile salmon started to increase in 1999, and forage fish densities increased in 2000, lagging by one year.     

Characterisation and preliminary quantification of the methane reservoir in a coastal sedimentary source: San Simón Bay,Ría de Vigo,NW Spain

Ría de Vigo is a river valley flooded by the sea, with a bay (San Simón Bay) at its innermost part. The accumulation of Holocene sediment in San Simón Bay has been studied by the integration of 1) large scale high resolution seismic data, and 2) detailed geochemical analysis of a gravity core. In San Simón Bay the majority of the seismic records are obscured by acoustic turbidity which represents gassy sediments, but on records from Rande Strait it is possible to distinguish two Quaternary seismic sequences; an Upper Pleistocene sequence (SQ1) and a Holocene sequence (SQ2). Only SQ2 is recognized in San Simón Bay where it is comprised of two seismic units; the upper unit represents the HST sediment, i.e. the period of highest sea level. A gravity core taken within the gassy zone at 10 m water depth provided 3.55 m of fine-grained sediments (muds) from the youngest seismic unit (4 m thick). Geochemical analysis show high values (4 to 10%) of TOC. Sediment and porewater analyses indicate a distinct sulphate–methane transition zone (SMTZ) between 60 and 80 cm where sulphate is depleted (to <1.7 mM) and methane increases (to >0.4 mM). The top of the acoustic turbidity (the gas front) at 80 cm corresponds to the lower limit of the SMTZ. The methane cannot have been derived from the underlying metamorphic and granitic rocks, but was probably derived by microbial degradation of the organic matter in the Holocene sediments. We estimate that the sediments of the Bay contain approximately 1.8 × 10⁶ m³ of organic carbon and 275 ton of methane.     

Abundance and distribution of ichthyoplankton in the upwelling zone off Oregon during anomalous El Niño conditions

The abundance and distribution patterns of nearshore ichthyoplankton were investigated during a year of anomalously high sea temperatures off Oregon. Samples collected from 2 to 18 km offshore from April through September of 1983 showed increased occurrences and higher abundances of taxa usually found at distances offshore of 37 km in other years. The dominant species collected, comprising more than half of the total larval fish abundance, was the northern anchovy (Engraulis mordax). Larval anchovy have rarely been collected inshore in previous studies. Many of the dominant taxa normally found inshore, especially osmerids, were present in reduced numbers in 1983. Changes in the hydrographic conditions associated with onshore surface drift and reduced summer upwelling during the 1983 El Niño could explain the distributional patterns observed. The warm inshore waters apparently provided a substantial spatial and temporal expansion of the spawning habitat for E. mordax.     

A 9-year time-series of planktonic foraminifer fluxes and environmental change in the Bering sea and the central subarctic Pacific Ocean, 1990–1999

A 9-year study of planktonic foraminifer fluxes was conducted in the Bering Sea (Station AB) and in the central subarctic Pacific (Station SA). Results clearly reflected variations of the water mass characteristics in the upper layers. The 9-year means of total foraminifer fluxes were the same (1400 shells m⁻² d⁻¹) at both stations. However, total foraminifer flux at Station AB tended to show its primary maximum during fall (October–December) and its secondary maximum in spring (April–June), whereas the primary maximum appeared in spring and the secondary maximum in fall at Station SA. Seasonal variation was more apparent at hemipelagic Station AB than at pelagic Station SA. Planktonic foraminifers found at both stations were of six species: Neogloboquadrina pachyderma, Globigerina umbilicata, Globigerinita glutinata, Globigerina quinqueloba, Globorotalia wilesi, and Orbulina universa. The foraminifer assemblages at the two stations reflected the temperature difference in the surface waters. The variable %G. umbilicata tended to be high in the warm surface waters during the summers. The temporal and geographical variation of %G. quinqueloba indicated that this taxon prefers regions with relatively low diatom fluxes. A notable appearance of O. universa occurred in 1997 at Station SA. During this period, other measured biogenic particle fluxes, such as those of diatoms, were low. This unusual 1997 event may be a reflection of global climatic change that happened to be observed in the central subarctic Pacific Ocean.     

The physical structure of a cold filament in a Chilean upwelling zone (Península de Mejillones, Chile, 23°S)

Cold filaments associated with Eastern Boundary Currents are typically narrower than 100 km but can be several hundred kilometers long, extending from the coast to the open ocean in upwelling areas. One such structure, observed off Península de Mejillones (23°S, Chile), was studied with both satellite images and two 5-days hydrographic cruises carried out during January 1997. The study used a coastal grid of 31 stations in an area of 165 ×155 km², approximately. The spatial distribution of the filament and its change between cruises are described from the horizontal distributions of dynamic height, temperature, salinity and dissolved oxygen. The filament was a shallow feature (thickness <100 m) and extended at least 165 km toward the open ocean. A meandering northward current flowed at the borders of the filament, separating oceanic and coastal waters of different physical properties. Comparisons of cross sections of the filament near the coast and in the oceanic zone show the ascent of the shallow salinity minimum (SSM), and its extension toward the ocean, bound to the filament. It is concluded that Subantarctic Water ((SAAW) distinguish by low salinity, high dissolved oxygen) and Equatorial Subsurface Water ((ESSW) high salinity, low dissolved oxygen, high nutrient content) form this filament, and that their relative proportions depend on the strength of the coastal upwelling. Thus, the knowledge of the dynamics of these structures is fundamental to better understanding of the spatial distribution of important biological variables, such as nutrients and chlorophyll, in the coastal ecosystem.