The shoaling of nutrient-enriched subsurface waters as a mechanism to sustain primary productivity off Central Baja California during El Niño winters

Using CalCOFI data for coastal shallow stations (above 100 m depth), higher than expected nitrate concentrations were detected in near-surface high-temperature waters off of Central Baja California during some El Niño winters. Though recent data are not available for Central Baja California, past El Niño data, though limited, showed nitrate concentrations above 16 μM at temperatures above 16 °C, and nitrate concentrations between 1 and 2 μM at 19 °C, while the previously established relationship of temperature and nitrate for California Current waters predicts nitrate depletion above 14 or 15 °C. The anomalous, high temperature–high nitrate enrichment events documented in Central Baja California were detected as shallow as 9 m and as deep as 73 m, were associated with low-oxygen (between 2 and 4 ml/l) and high-salinity (between 33.8 and 34.3 psu) waters, and occurred during the winter months of an El Niño year. Using recent data for San Diego, CA, similar but weaker enrichment events were detected for the El Niño winter of 1997–1998. The periodic shoaling of a subsurface subtropical water mass of high temperature, high salinity, low oxygen and high nutrients during some El Niño winters is proposed to cause periodic enrichment and to maintain productivity during warming events in this area. Enrichment events were not detected off Ensenada, in Northern Baja California, possibly due to the amplification of the onshore flow during El Niño there, or due to the Ensenada front. The proposed mechanism of periodic enrichment of nutrient-depleted surface waters during some El Niño winters by subsurface waters from the California Undercurrent may explain the following: (1) survival of giant kelp forests at their southern limit in Central Baja California documented during past El Niño events in warm waters, (2) the rapid recovery and high carrying capacity of giant kelp documented after the mass disappearance during El Niño 1997–1998, and (3) the increase in the extent of mesotrophic chlorophyll detected in the area during the 1997–1998 and 1982–1983 El Niño events.     

Dynamics and fishery of the Peruvian hake: Between nature and man

Mean length in the catch of Peruvian hake (Merluccius gayi peruanus) declined drastically by 8 cm in a few months in 1992. The 1991–93 El Niño event marked the beginning of changes in bottom environmental conditions, that along with other changes in the ecosystem, were the cause of the disappearance of large sized old hake from the traditional fishing grounds and the invasion of these areas by high concentrations of small sized young hake, traditionally distributed in the southern areas. A misinterpretation of this change gave an optimistic perception of the state of the stock, leading to a large increase of the fishing fleet and therefore to an increase of the fishing pressure on this resource. Catch increased quickly to very high levels in 1996, the fishery indicators of relative abundance remained stable, while global abundance of hake was diminishing. The resource responded to these natural variations of environment and to the intense fishing pressure with changes in some aspects of its biology and ecology. The most remarkable was the decrease of the length and age at first maturity that led to spawning by 1 year old (19 cm) individuals in a species whose observed longevity was 14 years.In 2002 a total and indefinite fishery closure was established by the Peruvian government and a technical commission for the hake recovery was created, which carried out continuous monitoring of the stock. After a closure of 20 months the indicators showed signs of recovery and the fishery was reopened in 2004. However, after an encouraging beginning during 2004 and the first quarter of 2005, neither abundance nor age structure had improved by the end of 2005, and in fact were deteriorating by the start of 2006. The survival of the age 2 and 3 groups was very low and few 4+ years old individuals were observed. Several factors working at different time scales have been proposed to explain the status of hake. The impact of the fishery, variations in the abiotic environment, predation by the jumbo squid (Dosidicus gigas) and cannibalism were considered as potential causal factors of this decline.     

Hydrographic structure and zooplankton abundance and diversity off Paita, northern Peru (1994 to 2004) — ENSO effects, trends and changes

The objective of the present study was to verify possible spatial, seasonal, and inter-annual changes in the zooplankton off Paita (northern Peru), an upwelling area located closely to the limits of cold Humboldt Current and warm Equatorial Surface Waters. Zooplankton was sampled at subsurface on 53 occasions from August 1994 to December 2004 at four stations located 2 to 30 km offshore with a WP-2 net (300 µm). Extremely high surface water temperatures combined with low salinities were observed during the 1997/98 El Niño up to 29.0 °C) and in April 2002 (up to 25.0 °C). Temperatures more than 2 °C above monthly average were also observed in October 1994, in April 2000, and in November 2004. Significant trends were observed for oxygen concentration (increase) and several horizontal and vertical gradients. Among the copepods (72% of all individuals), the most abundant species were Paracalanus parvus (28%), Acartia tonsa (26%), and Calanus sp. (10%). The strong 1997–98 El Niño (EN) event led to drastic changes in species composition that were reversed during the 1998–99 La Niña (LN) event. Community parameters such as total abundance, diversity, equitability and species richness displayed marked variations associated with the 1997–98 EN and long-term trends. Long-term trends were significant for several vertical and horizontal temperature and oxygen gradients, indicating an increase in upwelling intensity at the shelf during the study period. 10-year-trends were also significant for total zooplankton abundance (increase) and community evenness (J, decline). Our data confirmed the importance of the weak EN in 2002/03 for the study region. Within the trend of increasing zooplankton abundance, a sharp step or shift was observed from 1999 to 2000. When using sequential t-tests to detect shifts in (x + 1) transformed abundance data, a significant rupture was found between the last sampling in 1999 and the first sampling in 2000. Also, a substantial decrease in diel variability occurred after 1999, probably due to changes in vertical migration patterns. The considerable increase in zooplankton abundance over the study period, the ENSO effect, and the 1999–2000 transition are discussed with regard to synchronicity with other zooplankton time series. The present study contributes with the first evidence from an important area located in the Humboldt Current for synchronous trends and changes that were previously observed elsewhere in the Pacific. Our results demonstrated the importance of long-term zooplankton monitoring studies in upwelling areas, and confirms the idea of dramatic changes in pelagic ecosystem structure occurring in the East Pacific.     

Time-series analysis of remote-sensed chlorophyll and environmental factors in the Santa Monica–San Pedro Basin off Southern California

The time-series of remote-sensed surface chlorophyll concentration measured by SeaWiFS radiometer from September 1997 to December 2001 and the relevant hydrological and meteorological factors (remote-sensed sea surface temperature, atmospheric precipitation, air temperature and wind stress) in Santa Monica Bay and adjacent waters off southern California were analyzed using wavelet and cross-correlation statistical methods. All parameters exhibited evident seasonal patterns of variation. Wavelet analysis revealed salient long-term variations most evident in air temperature during El Niño 1997–1998 and in wind stress during La Niña 1998–1999. Short-period (<100 days) variations of remote-sensed chlorophyll biomass were mostly typical to spring seasons. Chlorophyll biomass was significantly correlated with air temperature and wind stress: an increase of chlorophyll biomass followed with 5–6-day time lag an increase of wind stress accompanied by a simultaneous decrease of air temperature. The mechanism of these variations was an intensification of phytoplankton growth resulting from the mixing of water column by wind stress and entrainment of nutrients into the euphotic layer.     

The influence of oceanographic scenarios on the population dynamics of demersal resources in the western Mediterranean: Hypothesis for hake and red shrimp off Balearic Islands

The aim of the present paper is to study the relationships between some climatic indices and parental stock, recruitment and accessibility to trawl fishery of hake (Merluccius merluccius) and red shrimp (Aristeus antennatus) off Balearic Islands (western Mediterranean). Available annual catch per unit effort, recruitment and spawning stock biomass have been used as biological data. As environmental data, the meso-scale IDEA index and the large-scale North Atlantic Oscillation (NAO) and Mediterranean Oscillation (MO) indices have been used. To analyze possible links between these indices with the population dynamics of demersal resources, two non-linear approaches have been applied: (i) stock–recruitment relationships from Ricker and Beverton–Holt models, by sequentially incorporating environment factors; (ii) generalized additive modelling, both classical general and threshold non-additive models were considered. The latter simulate an abrupt change in explicative variables across different phases (time periods or climatic index values). The results have shown that two oceanographic scenarios around the Balearic Islands, associated with macro and meso-scale climate regimes, can influence the population dynamics of hake and red shrimp. This is especially true for recruitment, which seems to be enhanced during low NAO and IDEA indices periods. During these periods, colder-than-normal winters generate high amounts of cold Western Mediterranean Intermediate Waters (WIW) in the Gulf of Lions, which flow southwards and reach the Balearic Islands channels in spring, increasing the productivity in the area. This oceanographic scenario could also be favourable to the distribution of hake on the fishing grounds where the trawl fleet targets this species, increasing its accessibility to the fishery. Both spawning stock and abundance of red shrimp seems to be also enhanced by high MO index periods, which could reflect the increased presence of the saline and warm Levantine Intermediate Waters (LIW) in the study area, extending over the fishing grounds of this species. The proposed interactions can be useful to assess and manage these important demersal resources.     

Variability in hydrographic conditions to the east and northwest of South Georgia, 1996–2001

Six years of high-resolution hydrographic data from the eastern and northwestern sides of South Georgia (southwest Atlantic) are used to study the changing circulation and water mass properties of the region. One year of data from these locations was used previously to describe the oceanographic conditions at those times; using the much greater volume of data now available, we identify which features appear temporally robust and which are transient, and begin addressing topics relating to the forcing of the inter-annual variability and the potential consequences for the local ecosystem. Waters on the shelf and those over the adjacent deep ocean invariably have different hydrographic properties, though the transition between them can be abrupt or gradual. The onshelf/offshelf differences vary greatly from year to year, due to the combined influences of local and remote processes. There are several instances of strong physical coupling between the eastern and northwestern sides of South Georgia; this offers potential for distinguishing physically-induced ecosystems changes separately from biologically-induced ecosystems changes. On the northeast side of the shelf, close to Cumberland Bay, there is evidence of an often intense, but variable, cyclonic circulation that is the result of interaction with the local bathymetry. This may act as a retention mechanism, and enhance local productivity. Two examples of extreme cold anomalies are present in the series of measurements. One of these (in 2000/2001) affected a limited area at the eastern side of the region surveyed and was due to an intrusion of the Southern Antarctic Circumpolar Current Front. The other (in early 1998) was due to the combined effects of the passage of a large-scale ocean anomaly that had its origins upstream in the Pacific Ocean, and strong air/sea interaction. Both of these were associated with the strong 1997/1998 El Niño event. Whilst previous studies have observed a link between El Niño forcing and ocean response around South Georgia with a temporal lag of around 3 years, we observe a much more rapid response to the extremely strong 1997/1998 El Niño event. This indicates that the ocean and ecosystem around South Georgia are more immediately susceptible to extreme instances of remote climatic forcing than had been supposed.     

Climate Variability and Estuarine Water Resources: A Case Study from Tampa Bay,Florida

Natural variability in the myriad of physical processes that impact and control estuaries occurs at time scales that typically may exceed or partly exceed many monitoring programs. With respect to documenting and monitoring impacts of human influences- on estuaries, it is therefore important to frame the results of short-term monitoring programs within the context of longer term natural variability in the environment. In the Tampa Bay, Florida area, climate variability strongly influences seasonal precipitation, stream flow, and salinity. In particular, El Niño-Southern Oscillation impacts during the winter have the potential to influence both the availability of surface water for water supply withdrawal and the evaluation of the impacts of surface water withdrawals on the ecosystem.     

An idealized model of interaction between fronds of the large seaweed Durvillaea antarctica

The effect of interactions between individual fronds in a bed of the large intertidal seaweed Durvillaea antarctica, when forced by breaking waves, is studied using a computational model. The model simulates the response of a seaweed bed using a sequence of connected oscillators which are excited by a propagating forcing function representing a breaking ocean wave. Two new facets of the interplay between seaweeds and hydrodynamics are considered: (i) wave forcing due to breaking waves is often not sinusoidal in the rocky intertidal zone and (ii) a frond interaction term is included. The addition of frond interaction reduces the maximum loading on individuals within the bed by around 30% using estimated biomechanical parameters. The rate of change of the loading on the holdfast (equivalent to the “jerk”) is affected in a similar or greater fashion.     

Environmental factors which affect growth of Japanese common squid, Todarodes pacificus, analyzed by a bioenergetics model coupled with a lower trophic ecosystem model

Bioenergetics model is applied to Japanese common squid, Todarodes pacificus. The temporal change of wet weight of common squid, which migrates in the Sea of Japan, is simulated. The time dependent horizontal distribution of prey is calculated a priori by 3-D coupled physical–biological model. The biological model NEMURO (North Pacific Ecosystem Model for Understanding Regional Oceanography) is used to simulate the lower-trophic ecosystem including three kinds of zooplankton biomass two of which is used as prey of common squid. A bioenergetics model reproduced appropriate growth curve of common squid, migrating in the North Pacific and the Sea of Japan. The results show that the wet weight of common squid in the northern Sea of Japan is heavier than that migrating in the central Sea of Japan, because prey density of the northern Sea of Japan is higher than that of the central Sea of Japan. We also investigate the wet weight anomaly for a global warming scenario. In this case, wet weight of common squid decreases because water temperature exceeds the optimum temperature for common squid. This result indicates that migration route and spawning area of common squid might change with global warming.     

The winter St. Helena climate index and extreme Benguela upwelling

Climate changes in the subtropical South-east Atlantic turn out to be well described by the St. Helena Island Climate Index (HIX) and observed fluctuations are in good agreement with inter-decadal variability of the entire South Atlantic Ocean. Year-to-year variations of the averaged austral winter HIX (July–September), representative of the main upwelling season, were compared with (i) corresponding averages of the geostrophic alongshore component of the south-east trade wind (SET) between St. Helena Island in the south-west and Luanda/Angola in the north-east, (ii) the meridional distribution of surface waters colder than 13 °C to characterise intense Benguela upwelling (IBU), and (iii) the meridional position of the Angola-Benguela Frontal Zone (ABFZ) determined by means of sea surface temperature images for offshore distances between 50 and 400 km. Temporal changes of these parameters were investigated and showed that the frequency of consecutive years of strong and relaxed Benguela upwelling is characterised by a quasi-cycle of about 11–14 years. It is proposed that the index of the winter HIX may be used as a ‘surveyor’s rod' to describe interannual changes in the Benguela upwelling regime as well as those of the embedded marine ecosystem.     

Ekman transport along the Galician Coast (NW, Spain) calculated from QuikSCAT winds

Ekman transport is studied close to the Galician coast by means of wind data provided by the QuikSCAT satellite from November 1999 to October 2005. Three different coastal zones are identified, western coast from Miño River to Cape Finisterre, middle coast from Cape Finisterre to Cape Ortegal and northern coast, from Cape Ortegal to Cape Peñas. In addition to existence of long-term variations, the periodicity of the transport signal is characterized by an annual component (365 days), a seasonal fluctuation (50–80 days) and a time scale related to passing storms (15–20 days). Although the periodicity of the signal is similar at the three zones due to external meteorological forcing, the Ekman transport is modulated by the presence of the coast, in such a way that seasonal patterns vary in intensity and direction along the coast. Thus, the spring–summer pattern is characterized by high transport at the western coast, pointing seaward perpendicular to the shore-line. The same orientation is observed at the middle coast although with a lower magnitude. Finally, Ekman transport at the northern coast points landward and oblique to the shore-line. The different transport orientations are shown to be responsible for the upwelling probability variation along the coast. On the other hand, the autumn–winter pattern does not show a clear trend with important inter-annual differences showing the high variability of Ekman transport for this period.     

Dynamics of suprabenthos-zooplankton communities around the Balearic Islands (western Mediterranean): Influence of environmental variables and effects on the biological cycle of Aristeus antennatus

Dynamics of suprabenthos and zooplankton were analyzed in two areas located in the NW (off Sóller harbour) and S (off Cabrera Archipelago) of Mallorca (Balearic Islands, western Mediterranean) at depths ranging between 135–780 m. Four stations situated respectively at 150 m (shelf-slope break), and at bathyal depths of 350, 650 and 750 m were sampled at bi-monthly intervals during six cruises performed between August 2003 and June 2004. Suprabenthos showed maximum biomass in both areas from late spring to summer (April to August), while minimum biomass was found in autumn (September–November). Though variable, temporal dynamics of zooplankton showed peaks of biomass in late winter and summer (February and June), while minimals occurred in autumn (August–September) and, at bathyal depths, in April. Suprabenthos (abundance; MDS analyses) showed a sample aggregation as a function of depth (3 groups corresponding to the shelf-slope break, upper slope — over 350 m; and the middle, deeper part of the slope — over 650–750 m), without any separation of hauls by season. By contrast, zooplankton samples were separated by season and not by depth. There was evidence of three seasonal groups corresponding to summer (June 2004–August 2003), autumn–winter (September and November 2003, February 2004), and spring (April 2004), being especially well established off Sóller. In general, suprabenthos was significantly correlated with the sediment variables (e.g. total organic matter content (% OM), potential REDOX), whereas zooplankton was almost exclusively dependent on Chl a at the surface, which suggests two different food sources for suprabenthos and zooplankton. The increase of suprabenthos abundance in April–June was paralleled by a sharp increase (ca. 2.8 times) in the %OM on sediment during the same period, coupled ca. 1–2 months of delay with the peak of surface Chl a recorded in February–March (from satellite imagery data). Suprabenthos biomass was also correlated with salinity close to the bottom, suggesting a link between suprabenthos abundance and changes in the oceanographic condition of water masses close to the bottom. It is suggested that a higher suprabenthos biomass recorded off Sóller in comparison to that off Cabrera in June could, in turn, be related to a seasonal inflow of Levantine Intermediate Water (LIW) in April–June in this area at mid bathyal depths (350–650 m). This trend would be based on: 1) it was evident only at mid-slope depths between 350–750 m, coinciding with the LIW distribution, and 2) it was not recorded among zooplankton (collected throughout the water column). The possible effect of the fluctuations of suprabenthos and zooplankton on higher trophic levels has been explored studying the diet and food consumption rates of the red shrimp Aristeus antennatus, as indicator species by its dominance in bathyal communities. A. antennatus increased its food consumption from February to April–June 2004 off Sóller, which in the case of large (CL > 40 mm) specimens was found in both areas. In addition, there was a shift of diet from winter to spring–early summer. In this last period, A. antennatus preyed upon euphausiids and mesopelagic decapods and fish, while benthos (e.g. polychaetes and bivalves) decreased in the diet. This indicates an increase in the food consumption and probably in the caloric content of the diet in pre-spawning females in April–June 2004, which is synchronized with the period when gonad development begins in A. antennatus females (May–June). Anyway, macrozooplankton, and not suprabenthos, is crucial as a high energetic food source in the coupling between food intake and reproduction in the red shrimp.     

Long-term sea surface temperature baselines—time series, spatial covariation and implications for biological processes

Coastal areas such as estuaries, bays and fjords usually have hydrographic characteristics (e.g., temperature, salinity) which differ from those at larger spatial scales and in offshore areas. The differences can arise if the areas are subject to different climatic forcing or if they are relatively isolated from each other due to topographic and ocean circulation features which inhibit advective inputs of water mass properties. Local differences in hydrographic conditions can therefore potentially limit the applicability of existing long time series of coastally monitored temperatures for addressing questions at large spatial scales, such as the response of species distributions and phenologies to climate change. In this study we investigate the spatial synchrony of long-term sea surface temperatures in the North Sea–Baltic Sea region as measured daily at four coastal sites (Marsdiep, Netherlands; Torungen, Norway; Skagens Reef, Denmark; and Christiansø, Denmark) and in several large offshore areas. All time series, including two series reconstructed and intercalibrated for this study (Skagens Reef and Christiansø, Denmark), began during 1861–1880 and continue until at least 2001. Temperatures at coastal sites co-varied strongly with each other and with opportunistically measured offshore temperatures despite separation distances between measuring locations of 20–1200 km. This covariance is probably due to the influence of large-scale atmospheric processes on regional temperatures and is consistent with the known correlation radius of atmospheric fluctuations (ca. 1000 km). Differences (e. g, long-term trends, amplitude of seasonal variations) between coastal temperatures and those measured in adjacent offshore areas varied nonrandomly over time and were often significantly autocorrelated up to 2 years. These differences suggest that spatial variations in physical oceanographic phenomena and sampling heterogeneities associated with opportunistic sampling could affect perceptions of biological responses to temperature fluctuations. The documentation that the coastally measured temperatures co-vary with those measured opportunistically in offshore areas suggests that the coastal data, which have been measured daily using standardized methods and instruments, contain much of the variability seen at larger spatial scales. We conclude that both types of time series can facilitate assessments of how species and ecosystems have responded to past temperature changes and how they may react to future temperature changes.     

Seasonal and short spatial patterns in European hake (Merluccius merluccius L.) recruitment process at the Balearic Islands (western Mediterranean): The role of environment on distribution and condition

This study evaluates the link between the recruitment process of European hake (Merluccius merluccius L.) of the Balearic Islands (western Mediterranean) and the environmental and physiological conditions. Spatio-temporal variation of abundance and condition of fish were evaluated at two locations each with different oceanographic conditions, one in the north (Sóller, SO) and another in the south (Cabrera, CA) of Mallorca Island. Environmental variables explored were hydrography, sediment characteristics, phytoplankton pigment concentration (ppc) and the trophic resources of hake. Individuals were divided in three life stages: recruits, post-recruits and young adults. Hepatosomatic index (HSI), relative condition index (K_n), gonadosomatic index (GSI) and fullness index (FI) were analysed for the three life stages. Recruitment starts in February with the incorporation of smaller hakes, and it can be followed through spring and early summer with a peak in April. However, some spatial heterogeneity in the recruitment process has been found between north and south of the Island. The main pulse of recruitment occurred at a different time in the two areas. Spatial heterogeneity was also consistent with the condition of hake recruits, with higher values of K_n and HSI at SO than at CA. Maximum values of K_n were found in February at SO and in April at CA, coinciding with the start of the different recruitment pulses to the fishing grounds. Post-recruits and young adults also showed higher condition at SO than at CA. The arrival in spring of the Western Winter Intermediate Waters (WIW) drives the spatial-temporal variation in abundance and condition of hake. Ppc was highly correlated with recruit abundance with a time lag of two months, while for post-recruits the time lag was three months. The observed differences in the condition of hake between areas could be a consequence of the fact that the waters to the north of Mallorca are comparatively more under the seasonal influence of WIW which is formed in more productive areas. Thus, this study characterises the short temporal and spatial variability in the hake recruitment process off the Balearic Islands, both in terms of abundance and fish condition. This pattern is explained on the basis of the mesoscale environmental variability observed between north and south of Mallorca and the ecological adaptive strategy of recruiting in the optimal environmental season.     

Trophic flows,ecosystem structure and fishing impacts in the South Catalan Sea,Northwestern Mediterranean

An exploited ecosystem from the continental shelf and upper slope of the Northwestern Mediterranean Sea was described by means of an Ecopath mass-balance model with the aim of characterising its functioning and structure and describing the ecosystem impacts of fishing. This application included some complexities added to the general modelling methodology due to the high biodiversity of the Mediterranean Sea and the multispecific nature of the fishery, and to the difficulties of working with fishing data which are usually irregularly or imprecisely collected. The model comprised 40 functional groups including primary producers, the main species of benthic, demersal and pelagic invertebrates, fishes and non-fish vertebrates and three detritus groups. In addition, trawling, purse seine, longline and troll bait fishing fleets were included.Results showed that the functional groups were organized into four trophic levels with the highest levels corresponding to anglerfish, dolphins, large pelagic fishes and adult hake. The system was dominated by the pelagic fraction, where sardine and anchovy prevailed in terms of fish biomasses and catches. Detritus and detritivorous groups also played key roles in the ecosystem and important coupled pelagic-demersal interactions were described. Considering Odum's theory of ecosystem development, the ecosystem was placed on an intermediate-low developmental stage due, at least partially, to the impact of fishing activity. This highlighted the high intensity of fishing in the ecosystem, in accordance with the general assessment of western Mediterranean marine resources, and fishing fleets were ranked as top predators of the system. The low trophic level of the catch was in line with the long history of exploitation in the area. However, the steady decline of pelagic landings between 1994 and 2003, coupled with a decrease of the pelagic biomass within the system, underlined the low resistance of the system in front of perturbations. This decline was reproduced under Ecosim dynamic simulations combining different scenarios of moderate increase of fishing effort and an environmental forcing affecting the availability of preys to small and medium-sized pelagic fishes under wasp-waist flow control.     

One-dimensional modelling of the plankton ecosystem of the north-western Corsican coastal area in relation to meteorological constraints

In order to study the influence of wind mixing on the spring variability of the plankton production of the north western Corsican coastal area, a one-dimensional (1D), vertical, coupled hydrodynamic/biological model (ECOHYDROMV) is used. A hydrodynamic 1D model of the water column with a k–l turbulent closure is applied. The biological model comprises six state variables, representing the plankton ecosystem in the spring period: phytoplankton, copepods, nitrate, ammonium, particulate organic matter of phytoplanktonic origin and particulate organic matter of zooplanktonic origin. The system is influenced by turbulence (expressed by the vertical eddy diffusivity), temperature and irradiance. The model takes into account momentum and heat surface fluxes computed from meteorological data in order to simulate a typical spring atmospheric forcing for the considered area. Results show that primary production vertical structure is characterised by a subsurface maximum which deepens with time and is regulated by the opposite gradients of nitrate concentration and irradiance. Surface plankton productivity is mainly controlled by turbulent vertical transport of nutrients into the mixed layer. The short time scale variability of turbulent mixing generated by the wind appears to be responsible for the plurimodal shape of plankton blooms, observed in the considered area. Furthermore, the model is applied to the study of the spring evolution of the plankton communities off the bay of Calvi (Corsica) for the years 1986 and 1988. In order to initiate and validate the model, time series of hydrological, chemical and biological data have been used. The model reproduces accurately the spring evolution of the phytoplankton biomass measured in situ and illustrates that its strong variability in those years was in close relation to the variability of the wind intensity.     

Winter phytoplankton blooms in the shallow mixed layer of the South China Sea enhanced by upwelling

The South China Sea (SCS) is the largest marginal sea in the world. Previous studies, including recent intensive paleo-oceanographic studies, suggest that the SCS is sensitive to many types of physical forcing on the short-term (e.g., internal waves and tides, mesoscale eddies, typhoons, etc.), annual (e.g., monsoon), inter-annual (e.g., El Niño), and very long-term (e.g., climate change) time scales. To better understand how various types of physical forcing influence biogeochemical cycles in the water column, a time-series study was initiated. Bimonthly hydrographic surveys occupied stations in the subtropical–tropic SCS at 19°N, 118.5°E. Results suggest that the Southeast Asian monsoons, northeasterly from October to April and southwesterly from May to September, have important effects on biogeochemical cycles in the upper water column. Hydrographic data showed that the mixed layer depth was much shallower in winter than in other seasons. During the winter monsoon period, the nitricline became shallower and upwelling sustained an elevated phytoplankton standing stock. Mean chlorophyll concentrations (0.65 mg Chl m^− 3) in winter were 8 times higher than in summer, and the integrated primary productivity over the euphotic zone reached as high as ca. 684 mg C m^− 2 day^− 1 in winter. The upwelling is produced by convergence of currents in the cyclonic gyre near the Luzon Strait, where the Kuroshio intrudes. In summer the current reverses following the wind change. The nitricline is depressed as downwelling occurs off northwest Luzon, resulting in strong nutrient limitation and very low chlorophyll concentrations.     

Impact of freshwater on a subarctic coastal ecosystem under seasonal sea ice (southeastern Hudson Bay, Canada). I. Interannual variability and predicted global warming influence on river plume dynamics and sea ice

Analysis of sea ice cover, runoff and air temperature observations in Hudson Bay shows marked interannual variability. This variability is thought to play a major role in determining overall productivity of the coastal ecosystem by changes to river plume extent, under-ice light conditions and nutrient levels during spring. Extensive field work off the Great Whale River in southeastern Hudson Bay has shown the importance of freshwater discharge, sea ice cover and meteorological forcing on the production of under-ice microalgae and the success of first feeding in fish larvae.Recent global climate model (GCM) results for a doubling of present atmospheric carbon dioxide indicate increases of both air temperature and precipitation in the Hudson Bay area. Predictions based on GCM results are used to estimate future changes to the sea ice and runoff regime. Sea ice breakup in the offshore is predicted to occur about one month earlier than presently. Estimates of the spring freshet in the Great Whale River indicate it will also advance by approximately one month. Onset of the spring freshet will occur about one month before Hudson Bay ice breakup, similar to present. A predicted reduction of about 35% in maximum sea ice thickness will lead to an increase in the ice-ocean interface irradiance and a decrease in melt water input to the Hudson Bay surface waters. These results are used in a discussion of potential effects of global climate change on northern coastal marine environments.     

Future aspects in marine ecosystem modelling

Existing ecosystem models are briefly presented and summarised. The problem of coupling physical and biological models as well as aspects of prediction and predictability are discussed. The general perception that marine ecosystems are inherently unpredictable due to non-linearity becomes questionable if the response of climate variability in marine ecosystems is analysed. Many authors have shown correlations between climate variability and the variability of abundance or biomass of marine organisms such as phytoplankton, zooplankton, benthos or fish recruitment in different parts of the world ocean. In the northern hemisphere, certain species show a linear response to climate variability mainly during winter and spring. However, the underlying mechanisms are not well understood. Often, a phase lag can be observed between climate variability and the reaction of organisms. The identification of a plausible mediator between climate and biology is difficult, since all possible physicochemical mechanisms having a direct or indirect influence on the variability of abundance or biomass of marine organisms have to be considered as mediator.The understanding of the reason of the phase lag, which possibly implies a “biological memory”, and the identification of all possible mediators are necessary to predict the response of marine organisms to climate variability. The identification of mediators will result in an improvement of coupled models, a deeper understanding of physical–biological interaction and the improvement of predictive capability of marine ecosystem models.     

Planktonic community structure and carbon cycling in the Arabian Sea as a result of monsoonal forcing: the application of a generic model

The Arabian Sea exhibits a complex pattern of biogeochemical and ecological dynamics, which vary both seasonally and spatially. These dynamics have been studied using a one-dimensional vertical hydrodynamic model coupled to a complex ecosystem model, simulating the annual cycle at three contrasting stations. These stations are characterised by seasonally upwelling, mixed-layer-deepening and a-seasonal oligotrophic conditions, respectively, and coincide with extensively measured stations on the two JGOFS ARABESQUE cruises in 1994. The model reproduces many spatial and temporal trends in production, biomass, physical and chemical properties, both qualitatively and quantitatively and so gives insight into the main mechanisms responsible for the biogeochemical and ecological complexity. Monsoonal systems are typified by classical food web dynamics, whilst intermonsoonal and oligotrophic systems are dominated by the microbial loop. The ecosystem model (ERSEM), developed for temperate regions, is found to be applicable to the Arabian Sea system with little reparameterisation. Differences in in-situ physical forcing are sufficient to recreate contrasting eutrophic and oligotrophic systems, although the lack of lateral terms are probably the greatest source of error in the model. Physics, nutrients, light and grazing are all shown to play a role in controlling production and community structure. Small-celled phytoplanktons are predicted to be dominant and sub-surface chlorophyll maxima are robust centers of production during intermonsoon periods. Analysis of carbon fluxes indicate that physically driven outgassing of CO₂ predominates in monsoonal upwelling systems but ecological activity may significantly moderate CO₂ outgassing in the Arabian Sea interior.