Validation of the 3D biogeochemical model MIRO&CO with field nutrient and phytoplankton data and MERIS-derived surface chlorophyll a images

This paper presents results obtained with MIRO&CO-3D, a biogeochemical model dedicated to the study of eutrophication and applied to the Channel and Southern Bight of the North Sea (48.5°N–52.5°N). The model results from coupling of the COHERENS-3D hydrodynamic model and the biogeochemical model MIRO, which was previously calibrated in a multi-box implementation. MIRO&CO-3D is run to simulate the annual cycle of inorganic and organic carbon and nutrients (nitrogen, phosphorus and silica), phytoplankton (diatoms, nanoflagellates and Phaeocystis), bacteria and zooplankton (microzooplankton and copepods) with realistic forcing (meteorological conditions and river loads) for the period 1991–2003. Model validation is first shown by comparing time series of model concentrations of nutrients, chlorophyll a, diatom and Phaeocystis with in situ data from station 330 (51°26.00′N, 2°48.50′E) located in the centre of the Belgian coastal zone. This comparison shows the model's ability to represent the seasonal dynamics of nutrients and phytoplankton in Belgian waters. However the model fails to simulate correctly the dissolved silica cycle, especially during the beginning of spring, due to the late onset (in the model) of the early spring diatom bloom. As a general trend the chlorophyll a spring maximum is underestimated in simulations. A comparison between the seasonal average of surface winter nutrients and spring chlorophyll a concentrations simulated with in situ data for different stations is used to assess the accuracy of the simulated spatial distribution. At a seasonal scale, the spatial distribution of surface winter nutrients is in general well reproduced by the model with nevertheless a small overestimation for a few stations close to the Rhine/Meuse mouth and a tendency to underestimation in the coastal zone from Belgium to France. PO₄ was simulated best; silica was simulated with less success. Spring chlorophyll a concentration is in general underestimated by the model. The accuracy of the simulated phytoplankton spatial distribution is further evaluated by comparing simulated surface chlorophyll a with that derived from the satellite sensor MERIS for the year 2003. Reasonable agreement is found between simulated and satellite-derived regions of high chlorophyll a with nevertheless discrepancies close to the boundaries.     

Vertical profiles of particulate organic matter and its relationship with chlorophyll-a in the upper layer of the NE Mediterranean Sea

Particulate organic matter (POM), nutrients, chlorophyll-a (CHL) and primary production measurements were performed in the upper layer of three different regions (cyclonic, anticyclonic and frontal+peripherial) of the NE Mediterranean Sea in 1991–1994. Depth profiles of bulk POM exhibited a subsurface maximum, coinciding with the deep chlorophyll maximum (DCM) established near the base of the euphotic zone of the Rhodes cyclone and its periphery, where the nutricline was situated just below the euphotic zone for most of the year. Moreover, the POM peaks were broader and situated at shallower depths in late winter–early spring as compared to its position in the summer–autumn period. Under prolonged winter conditions, as experienced in March 1992, the characteristic POM feature disappeared in the center of the Rhodes cyclone, where the upper layer was entirely occupied by nutrient-rich Levantine deep water. Deep convective processes in the cyclonic gyre led to the formation of vertically uniform POM profiles with low concentrations of particulate organic carbon (POC) (2.1 μM), nitrogen (0.21 μM), total particulate phosphorus (PP) (0.02 μM) and chlorophyll-a (0.5 μg/L) in the euphotic zone. Though the Levantine deep waters ascended up to the surface layer with the nitrate/phosphate molar ratios (28–29) in March 1992, the N/P molar ratio of bulk POM in the upper layer was low as 10–12, indicating luxury consumption of phosphate during algal production. Depth-integrated primary production in the euphotic zone ranged from 38.5 for oligotrophic autumn to 457 mg C m⁻² day⁻¹ for moderately mesotrophic cool winter conditions.     

Annual cycle of distribution of three ice-associated copepods along the coast near Dumont d'Urville, Terre Adélie (Antarctica)

In polar regions sea ice is a site of enhanced primary production during winter and provides important habitat for small grazers, such as copepods. We sampled zooplankton from the sea ice and water column throughout 2005, near Dumont d'Urville station (Terre Adélie, Antarctica). Three species of ice-associated copepods were found: two calanoid copepods Paralabidocera antarctica and Stephos longipes and the harpacticoid copepod Drescheriella glacialis. P. antarctica was the most abundant of the three and was closely associated with the sea ice during most of the year. This species had a one year life cycle with a probable over-wintering period in the sea ice as nauplii and a short copepodite phase in spring. Reproduction and spawning occurred in early summer. A comparison with two other populations (near Syowa and Davis stations) along the east coast of Antarctica showed that there was a temporal shift in the life cycles of the three populations, which was linked to variability in sea ice conditions. D. glacialis was the second most abundant copepod and was more common during the winter than during summer, indicating its preference for the sea ice habitat. In autumn, the presence of D. glacialis in the deeper part of the water column suggested that this species colonised the sea ice from the benthos. S. longipes was found only in the water column near Dumont d'Urville and was not very abundant. In Terre Adélie particular environmental conditions, such as the absence of a permanent sea ice zone throughout the year, a longer time of open water, strong katabatic winds and the presence of polynyas, have influenced both the abundance and distribution of the three common ice-associated copepods.     

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.     

Egg production of the copepod Acartia hongi in Kyeonggi Bay, Korea

Many studies of copepod egg production have shown that food availability and temperature are major factors that influence copepod growth. However, coastal environments are complicated ecosystem and the relationships between growth of copepods and influencing factors are not always clear in nature. We conducted a study along an inner–middle–outer bay transect where variations in environmental parameters would be expected to affect the biomass and egg production rate of A. hongi from February 2001 to December 2001. In this study, we investigated the abundance and biomass with developmental stages and egg production rates of A. hongi in relation to various environmental factors. The copepod A. hongi occurred continuously throughout the year, with a peak abundance in May. In general, the variation in egg production rates showed a similar tendency with the variations in chlorophyll-a throughout the study period. This suggests that phytoplankton biomass is an important factor that affects the egg production of A. hongi. In addition, during the warm season, the egg production of A. hongi was also influenced by the ciliates abundance in the middle and outer bay. Consequently, the egg production of A. hongi is generally affected by food availability in Kyeonggi Bay.     

Seasonal variability of the Black Sea chlorophyll-a concentration

The optimal spectral decomposition (OSD) method is used to reconstruct seasonal variability of the Black Sea horizontally averaged chlorophyll-a concentration from data collected during the NATO SfP-971818 Black Sea Project in 1980–1995. During the reconstruction, quality control is conducted to reduce errors caused by measurement accuracy, sampling strategy, and irregular data distribution in space and time. A bi-modal structure with winter/spring (February–March) and fall (September–October) blooms is uniquely detected and accurately documented. The chlorophyll-a enriched zone rises to 15 m depth in winter and June, and deepens to 40 m in April and 35 m in August. The June rise of the chlorophyll-a enriched zone is accompanying by near-continuous reduction of upper layer maximum chlorophyll-a concentration.     

Production of Acartia omorii (Copepoda: Calanoida) in Ilkwang Bay, southeastern coast of Korea

Production of the marine calanoid copepod Acartia omorii was measured from 2 October 1991 to 8 October 1992 at a station in Ilkwang Bay on the southeastern coast of Korea. A. omorii (nauplii + copepodites + adults) were present in the plankton throughout the year, with seasonal variation in abundance. Biomass of A. omorii was averaged at 0.44 mgC m^− 3, with peaks in February and July, and relatively low biomass in late summer and fall. Egg production rate ranged from 2.4 to 151.9 μgC m^− 3 day^− 1, which was equivalent to 95–6075 eggs m^− 3 day^− 1. Fecundity of an adult female was averaged at 38 eggs female^− 1 day^− 1. Instantaneous growth rates of copepodites were higher than those of nauplii stages. Annual production of A. omorii ranged from 33.5 mgC m^− 3 year^− 1 to 221 mgC m^− 2 year^− 1, showing a seasonal variation of daily production rate with peaks in February and July. The daily production rate of A. omorii was significantly correlated with chlorophyll a concentration. These results suggest that standing stocks and/or productivity of phytoplankton are the major influencing factors, rather than water temperature for the seasonal variation of production of A. omorii in Ilkwang Bay.     

Distribution of phytoplankton in the southern Black Sea in summer 1996, spring and autumn 1998

The species composition, abundance, and biomass of micro- (>15 μm) and nano- (<15 μm) phytoplankton were studied along the southern Black Sea during June–July 1996 and March–April and September 1998. A total of 150 species were identified, 50% of them being dinoflagellates. The average total phytoplankton abundance changed from 77×10³ cells l⁻¹ in spring to 110×10³ cells l⁻¹ in autumn and biomass from 250 μg l⁻¹ in summer to 1370 μg l⁻¹ in spring. Based on the extensive sampling grid from June–July 1996, phytoplankton seemed to have a rather homogeneous biomass distribution in the southern Black Sea. In all periods, the coccolithophorid Emiliania huxleyi was the most abundant species, its contribution to the total abundance ranging from 73% in autumn to 43% in spring. However, in terms of biomass, diatoms made up the bulk of phytoplankton in spring (97%, majority being Proboscia alata) and autumn (73%, majority being Pseudosolenia calcar-avis), and dinoflagellates in summer (74%, Gymnodinium sp.). There was a remarkable similarity in the dominant species between the western and eastern regions of the southern Black Sea, indicating transport of phytoplankton within the basin.     

Using satellite observations of ocean color to categorize the dispersal patterns of river-borne substances in the Gaoping (Kaoping) River, Shelf and Canyon system

Mapping the water constituents from remotely sensed ocean color data enables a better understanding of the dispersal patterns of river-borne substances in the Gaoping (formerly spelled Kaoping) River, Shelf and Canyon (KPRSC) system. Based on twelve MODIS-Aqua images in the KPRSC region taken in 2005, we apply a newly developed GA-SA approach to derive maps of chlorophyll-a concentration (Chl-a), colored dissolved organic matter (CDOM) and non-algal particle/detritus/mineral (NAP). The results demonstrated that the different characteristics of Chl-a, CDOM and NAP make them ideal tracers for observing large-scale dispersal patterns. With ancillary information of averaged daily precipitation, the daily wind field obtained from QuikSCAT (Quick Scatterometer), and the 8-day composite of the temperature field obtained from MODIS-Aqua, we categorized the surface dispersal patterns as coastal, northwestward and frontal patterns. Also, for the first time, we observed a sudden increase of biomass on a large scale from a pair of ocean color images taken over only a 2-day interval. Another remarkable feature is the interaction between the southeastward flow and the intrusion of the Kuroshio Branch, resulting in complicated patterns with various scales of vortex structures and current fronts. The observed features could be used for model validation of the flow field of the KPRSC system.     

Modelling the effects of physical–biological interactions and spatial variability in spawning and nursery areas on transport and retention of sardine Sardinops sagax eggs and larvae in the southern Benguela ecosystem

A particle-tracking individual-based model (IBM) coupled with a 3-D hydrodynamic model was used to investigate how spatial variability in spawning and nursery grounds may influence transport and retention of sardine (Sardinops sagax) eggs and larvae in the southern Benguela ecosystem. A temperature-dependent Bělehrádek development model based on laboratory growth data was used to determine the duration of one egg and two larval stages. Successful transport/retention depended on each particle's stage of development rather than its age. Results show that recruitment could occur in two distinct nursery areas on the west and south coasts of South Africa. Three viable recruitment ‘systems’ were identified: two are retention-based (spawned and retained on the west coast (WC–WC) or the south coast (SC–SC)) and one is transport-based (spawned on the south coast and transported to the west coast (SC–WC)). In the WC–WC system, the vertical distribution of eggs influenced retention; at intermediate depths of spawning (25–50 m) eggs avoided both offshore Ekman drift and deep cold water. In the SC–SC system, the area of spawning was important; > 50% of eggs from the eastern Agulhas Bank (EAB) were retained in the south coast nursery grounds, whereas very few eggs were transported there from west of Cape Agulhas. In the SC–WC system, area of spawning was also important; 40% of the eggs spawned on the western Agulhas Bank (WAB) were transported to the west coast nursery ground. Sardine life history strategy could be divided between two main systems: the west coast system (spawning on the WAB and WC, and recruiting to the WC) and the Agulhas Bank system (spawning on the central and eastern Agulhas Bank, and recruiting to the SC).     

Decadal-scale variations of trophic levels at high trophic levels in the Yellow Sea and the Bohai Sea ecosystem

A total of 2759 stomachs collected from a bottom trawl survey carried out by R/V “Bei Dou” in the Yellow Sea between 32°00 and 36°30N in autumn 2000 and spring 2001 were examined. The trophic levels (TL) of eight dominant fish species were calculated based on stomach contents, and trophic levels of 17 dominant species in the Yellow Sea and the Bohai Sea reported in later 1950s and mid-1980s were estimated so as to be comparable. The results indicated that the mean trophic level at high trophic levels declined from 4.06 in 1959–1960 to 3.41 in 1998–1999, or 0.16–0.19·decade^− 1 (mean 0.17·decade^− 1) in the Bohai Sea, and from 3.61 in 1985–1986 to 3.40 in 2000–2001, or 0.14·decade^− 1 in the Yellow Sea; all higher than global trend. The dominant species composition in the Yellow Sea and the Bohai Sea changed, with the percentage of planktivorous species increases and piscivorous or omnivorous species decreases, and this was one of the main reasons for the decline in mean trophic level at high tropic levels. Another main reason was intraspecific changes in TL. Similarly, many factors caused decline of trophic levels in the dominant fish species in the Yellow Sea and the Bohai Sea. Firstly, TL of the same prey got lower, and anchovy (Engraulis japonicus) as prey was most representative. Secondly, TLs of diet composition getting lower resulted in not only decline of trophic levels but also changed feeding habits of some species, such as spotted velvetfish (Erisphex pottii) and Trichiurus muticus in the Yellow Sea. Thirdly, species size getting smaller also resulted in not only decline of trophic levels but also changed feeding habits of some species, such as Bambay duck (Harpodon nehereus) and largehead hairtail (Trichiurus haumela). Furthermore, fishing pressure and climate change may be interfering to cause fishing down the food web in the China coastal ocean.     

Environmental causes of the fluctuations of red shrimp (Aristeus antennatus) landings in the Catalan Sea

The fluctuations of catches of the deep-sea shrimp Aristeus antennatus (Risso, 1816) were analysed in the 6 ports of Catalonia where production is concentrated, based on monthly landings from 1988 to 2004. The 6 selected ports produced 300 t of red shrimp in 2004, or 80% of the total production in Catalonia. The series for each port showed clear interannual variability, with peaks of production in the early 1990s and more recently from 2001 to 2003. The time periods of the monthly data series, estimated by frequency analysis based on Fourier transform, varied around 7–8 years in the four central ports and 12–13 years in the two northern ports. Additionally, the different curves were not in phase: even in nearby ports, the maximum production is observed in different years. Since the North Atlantic Oscillation (NAO) index is an excellent proxy for long-term series of environmental variables, we aimed to explore relationships between the fluctuation of the NAO index and A. antennatus landings in the Catalan Sea. The correlation between the mean annual NAO index and the annual catches in each port was positive and significant with some time lags (from 1 to 3 years). The existence of clear patterns linking the NAO with marine ecological processes has been demonstrated in many studies, but the underlying ecological mechanisms are far from being well understood. The variations in environmental parameters linked to the NAO may act on biological organisms at different levels (individual, population) through physiology (metabolic and reproductive processes) or through trophic relationships, including ecological cascade effects. We propose that NAO-induced environmental variability may enhance food supply to A. antennatus and hence strengthen the reproductive potential of particular year classes, which result in increased catches 1 to 3 years later, although other possible effects of environmental variability on the population dynamics of this species are worth investigating.     

Role of filtering and biodeposition by Adamussium colbecki in circulation of organic matter in Terra Nova Bay (Ross Sea, Antarctica)

At Terra Nova Bay, the scallop Adamussium colbecki (Smith, 1902) characterises the soft and hard bottoms from 20 to 80 m depth, constituting large beds and reaching high values of density (50–60 individuals/m²) and biomass (120 g/m² DW soft tissues). To assess its role in the organic matter recycling in the coastal ecosystem, its filtering and biodeposition rates were evaluated in laboratory experiments during the austral summer 1993/94. Filtration rates, measured in a flow-through system, were calculated from the difference in particulate organic carbon (POC), nitrogen (PON) and chlorophyll-a (Chl-a) concentration in inflow and outflow water. Experiments were performed using natural sea water with POC, PON and Chl-a concentrations of about 450 μg/l, 90 μg/l and 2 μg/l, respectively. The biodeposition rate and the biochemical composition of the biodeposits were studied in order to detect how the organic matter is transformed through feeding activity of A. colbecki. At +1°C temperature, the average filtering rate was about 1 l h⁻¹ g⁻¹ (DW soft tissues) in specimens ranging in body mass from 2 to 3 g (DW soft tissues) and 6–7 cm long. The biodeposition rate in 3–8 cm long specimens, ranging from 0.4 to 5.7 g (DW soft tissues), was about 5.65 mg DW/g DW/day, leading to an estimate of Corg flux, through biodeposition by A. colbecki, of about 21 mg C m⁻² day⁻¹ at in situ conditions. Comparison between the biochemical composition of seston and biodeposits shows a decrease of the labile compounds, of the Chl-a/phaeopigments ratio in the biodeposits. The recorded C/N ratio decrease suggests a microbial colonisation in the biodeposits. This study suggests that Adamussium colbecki plays an important role in coupling the material fluxes from the water column to the sea bed, processing about 14% of total Carbon flux from the water column to the sediments, with an assimilation efficiency of 36%.     

Pigments, size and distribution of Synechococcus spp. in the Black Sea

Pigments, size and distribution of Phycoerythrin-containing unicellular cyanobacteria Synechococcus spp. within the euphotic zone were studied for the first time in April–May 1994 in the western and southwestern Black Sea by epifluorescence microscopy and flow-cytometry. Synechococcus was present in varying quantities at every station and depth studied. Surface spatial distribution of Synechococcus revealed that cells were much more abundant in offshore waters than near coastal regions under the direct influence of the Danube river. Minimum and maximum cell concentrations ranged between 9×10² and 1.45×10⁵ cells/ml at the surface, between 2×10³ and 1.23×10⁵ cells/ml at the chlorophyll sub-maximum layer, and between 1.3×10² and 3.5×10² at the nitrite maximum layer. Cells at the chlorophyll sub-maximum layer (based on in-situ fluorometer readings) fluoresce brighter and longer than the ones at the surface and lower depths. Spectral properties of chromophore pigment types of total 64 clonal isolates from different depths down to the lower layer of the euphotic zone (60 m) in the southern Black Sea coast revealed that all have type 2 phycoerythrobilin in common, lacking in phycourobilin. In vivo fluorescence emission maxima for the phycoerythrobilin were about the same (578 nm) for all isolates. All isolates examined showed in vivo absorption maxima at between 435 and 442 nm and at about 681 nm due to chlorophyll-a. Based on the flow cytometer mean forward light scatter data for size distribution, it could be concluded that cells at the surface mixed layer (0–10 m) were larger in cell size than the cells at lower depths (20–60 m).     

Evidence of episodic changes in deep-sea mesozooplankton abundance and composition in the Levantine Sea (Eastern Mediterranean)

Mesozooplankton samples from the 4270-m deep Ierapetra Basin in the oligotrophic Levantine Sea have revealed a strong temporal variability in abundance and composition hitherto unknown for the deep sea pelagic zone. The phenomenon reflected by a survey in June 1993 as compared to January 1987 is assessed by (i) the increase of the mesozooplankton standing crop throughout the water column, (ii) the absence of a decline in mesozooplankton abundance with increasing depth below 1000 m, (iii) the outstanding dominance of two interzonal calanoid copepods at bathypelagic and abyssopelagic depths and (iv) a substantial faunal change due to the codominance of Calanus helgolandicus, which was found for the first time in the area. The underlying hydrographical factors are addressed in the light of the Eastern Mediterranean Transient (EMT). Apart from a possible long-term faunal change, the EMT may exaggerate episodic plankton blooms and surface-abyssopelagic coupling in space and time.     

Observations on colony formation by the cosmopolitan phytoplankton genus Phaeocystis

Few marine phytoplankton have heteromorphic life cycles and also often dominate the ecosystems in which they occur. The class Prymnesiophyceae contains a notable exception: the genus Phaeocystis includes three species that form gelatinous colonies but also occur within their ranges as solitary cells. Phaeocystis antarctica and P. pouchetii are exclusively high latitude taxa, and are notable for regionally tremendous blooms of the colony stage. P. globosa occurs circumglobally, yet its colony blooms primarily are confined to colder waters within its range. Three additional species are warm water forms that have been reported only as solitary cells or loose aggregations that bear little resemblance to the organized colonies of the other taxa. Interpretation of existing data indicates that resource availability (light, temperature and nutrients) by itself is not sufficient to explain this distinction between cold-water colony-forming taxa and warm water solitary cell taxa, nor why colony development in P. globosa is essentially a spatially restricted phenomenon within a much broader geographic range. Colony development by P. globosa in situ has been observed at temperatures ≥20 °C, but only rarely and generally under conditions of seasonally or anthropogenically elevated nutrient supply. Data presented here demonstrate colony development at 20–22 °C in natural plankton communities from oligotrophic waters that were pre-screened through 63 μm mesh (i.e. lacking mesozooplankton and large microzooplankton), but not in unscreened communities containing microzooplankton and >63 μm zooplankton. Reduction of colony proliferation at higher temperatures by mesozooplankton grazing remains as an intriguing possibility that is consistent with available evidence to help explain differences in latitudinal extent of in situ colony development. These data are interpreted within a theoretical framework regarding the potential advantages and disadvantages of the two life cycle stages.     

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.     

Small scale retentive structures and Dinophysis

Despite its rarity, Dinophysis acuminata is in terms of economic impact, the first toxic algal species along the coasts of Western Europe. It is observed at low levels (< 20 cell l^− 1) all the year round but toxic events occur mainly in late spring and summer. D. acuminata ecophysiology is largely unknown due to the inability to culture it. Therefore, standard biomass models based on inorganic nutrition are largely inadequate. Presently, any progress in describing the conditions of population growth of this species will be a step forward to prediction of harmful events at the coast. This species has been observed at increased, albeit low cell densities in retentive eddies located in pycnocline layers. A concentration build-up of one species results from the balance between growth and loss processes, one of the latter being dispersal. The scales of interest for a D. acuminata population are of the order of 10 nautical miles on the horizontal and duration of 10 days, for a reported achievable growth rate of 0.6 day^− 1. A three dimensional (3D) hydrodynamical model of the Bay of Biscay has been elaborated to reproduce hydrological structures over the last decade. We attempt here to relate the existence of retentive structures revealed from simulations under realistic forcing conditions and the toxic coastal events recorded in the 10-year time series of the French plankton monitoring network database. The eddies in the coastal area appear to be directly related with the Dinophysis coastal events and they may be a potential effective tool to predict those.