Climate–ocean variability and Pacific hake: A geostatistical modeling approach

Climate forcing of the California Current has been known to impact the distribution and abundance of a number of local fish populations, but the mechanisms involved remain poorly understood. Climate metrics such as the Pacific Decadal Oscillation (PDO) and the El Niño Southern Oscillation (ENSO) are usually used to represent climate processes and direct links are made between climate forcing and production variability. This involves aggregation of impacts across large spatial scales and range of species. However, fluctuations in productivity are often the result of changes in physical habitat. In order to fully understand the relationship between climate and productivity, habitat changes should be addressed. In this study we use a geostatistical approach to quantify adult Pacific hake habitat during different climate regimes. Several authors have suggested that the distribution and intensity of the sub-surface poleward flow (the undercurrent) plays a key role in defining adult hake habitat along the west coast of North America. Here we build a model designed to predict hake habitat distribution in space based on sub-surface poleward flow distribution and bottom depth. Our results show that hake habitat expands in 1998 El Niño year compared to 1995. Given the important predatory role that hake plays in the CC, the amount and distribution of adult hake habitat has large implications for the Pacific Northwest food web and could thus serve as an ecosystem indicator representing important physical–biological interactions. Spatially based ecosystem indicators such as the one we develop here address two important yet neglected areas in the ‘Ecosystem Indicators debate’: the importance of developing metrics explicitly representing spatial and environmental processes shaping ecosystem structure. Without these, our power to fully describe ecosystems will be limited.     

Gulf of Gdansk spring bloom physical, bio-optical, biological modelling and contact data assimilation

In this study we propose a model of phytoplankton population dynamics in the marine ecosystem, which includes physical, biological and bio-optical parts. As an example we simulate the abnormal 1993 Gulf of Gdansk spring bloom, when extremely high chlorophyll concentrations were observed. For the one-dimensional model we use two different methods of contact chlorophyll observation assimilation to fit a model of “in situ” data. The results are compared with two-dimensional ecosystem modelling based on a barotropic model of wind-driven circulation without assimilation.     

Integrating Climate and Ocean Change Vulnerability into Conservation Planning

Tropical coastal and marine ecosystems are particularly vulnerable to ocean warming, ocean acidification, and sea-level rise. Yet these projected climate and ocean change impacts are rarely considered in conservation planning due to the lack of guidance on how existing climate and ocean change models, tools, and data can be applied. Here, we address this gap by describing how conservation planning can use available tools and data for assessing the vulnerability of tropical marine ecosystems to key climate threats. Additionally, we identify limitations of existing tools and provide recommendations for future research to improve integration of climate and ocean change information and conservation planning. Such information is critical for developing a conservation response that adequately protects these ecosystems and dependent coastal communities in the face of climate and ocean change.     

Spatial and temporal variability in the pelagic ecosystem of the East Sea (Sea of Japan): A review

The East Sea (Sea of Japan) is a unique marginal sea because it exhibits features of oceanic dynamics of much larger ocean basins. This semi-enclosed basin may be considered as a model or microcosm for understanding of how biological processes and distributions in pelagic ecosystem are interacting with physical processes in highly dynamic ocean regions. This overview summarizes the recent progresses concerning spatial and temporal variability of pelagic ecosystem components form an interdisciplinary point of view. Spatial characteristics of physical environments and biogeography in the region are distinguished mainly by the subpolar front. It was also found that long-term changes in biomass and community structure as well as those in the physical and biological environments are associated with climate variability in the region. We conclude by identifying main needs for the information and researches, particularly regular and long-term sampling, and permanent monitoring if possible.     

Topological constraints on the dynamics of wasp-waist ecosystems

Small pelagic fish species like anchovy or sardines are of high ecological and economical importance. As marine food webs are fished down, these small pelagics tend to be more exploited and overfished. It is not yet very well known what the possible effects of their collapse can be, therefore there is an urgent need to outline a theoretical framework for understanding their dynamics. These fish occupy very special position in food webs, ensuring energy transfer between species lower and higher levels, while forming narrow “wasp-waists” poor in number of species (but very abundant). Our purpose was to quantify the interaction structure of model food webs of equal complexity but different levels of “wasp-waistedness”. We analysed the topological properties of the webs by characterising every direct and indirect interactions between individual species, as well as by assessing the relative positional importance of each species in each web. We found that (1) the shorter the interaction pathways considered, the weaker the predictive power of node degree for positional importance, (2) the importance of species varies more in wasp-waist food webs, (3) if longer indirect chain effects are considered, indirect effects can well be stronger than direct ones, (4) interactions between coexisting wasp-waist species are stronger than the average, and (5) the “self-regulatory” looping effects are also stronger for wasp-waist species. Based on the topological properties of the networks, our results describe constraints acting on the dynamical behaviour of wasp-waist ecosystems. We give explanations, from this viewpoint, for regime shifts in which one WW species replaces another, and for the unpredictable dynamics of these fish stocks. From a marine conservation viewpoint, we illustrate that as the abundance of wasp-waist species decreases, the architecture of energy flows becomes highly vulnerable and unreliable. We provide an approach for quantifying these structural changes.     

A generalized model of pelagic biogeochemistry for the global ocean ecosystem. Part I: Theory

The set of equations for global ocean biogeochemistry deterministic models have been formulated in a comprehensive and unified form in order to use them in numerical simulations of the marine ecosystem for climate change studies (PELAGOS, PELAgic biogeochemistry for Global Ocean Simulations). The fundamental approach stems from the representation of marine trophic interactions and major biogeochemical cycles introduced in the European Regional Seas Ecosystem Model (ERSEM). Our theoretical formulation revisits and generalizes the stoichiometric approach of ERSEM by defining the state variables as Chemical Functional Families (CFF). CFFs are further subdivided into living, non-living and inorganic components. Living CFFs are the basis for the definition of Living Functional Groups, the biomass-based functional prototype of the real organisms. Both CFFs and LFGs are theoretical constructs which allow us to relate measurable properties of marine biogeochemistry to the state variables used in deterministic models. This approach is sufficiently generic that may be used to describe other existing biomass-based ecosystem model.     

Effect of mixing on microbial communities in the Rhone River plume

The biological processes involved during mixing of a river plume with the marine underlying water were studied off the Rhone River outlet. Samples of suspended and dissolved matter were collected while tracking a drifting buoy. Three trajectories were performed, at 2-day intervals, under different hydrological and meteorological situations. A biological uptake was evidenced from ammonium (NH₄) and phosphate (PO₄) shortage, indicating an early “NH₄-dependent” functioning occurring before the well-known “NO₃-based” cycle. The different ratios between NH₄, NO₃ and PO₄, as a function of salinity, were discussed to detail the preferential use in PO₄ and NH₄. Salinity zones with enhanced bacterial production, high chlorophyll a concentration, as well as DOC, NH₄ and PO₄ consumption were evidenced from 20 to 35 in salinity. It was shown that the successive abundance of bacteria and phytoplankton during transfer reflected the competition for PO₄ of both communities. On the Rhone River plume, the role played by temperature, light conditions and suspended matter upon biological activity seems relatively minor compared to salinity distribution and its related parameter: nutrient availability. It can be concluded that biological uptake in the Rhone River plume was closely related to the dilution mechanism, controlled itself by the dynamics of the plume. In windless conditions and close to the river mouth, the density gradient between marine and river water induced limited exchanges between the nutrient-rich freshwater and the potential consumers in the underlying marine water. Consequently, little biological activity is observed close to the river mouth. Offshore, mixing is enhanced and a balance is reached between salinity tolerance and nutrient availability to form a favourable zone for marine phytoplankton development. This can be quite far from the river mouth in case of a widely spread plume, corresponding to high river discharge. Under windy and wavy conditions, the plume freshwater is early and rapidly mixed, so that the extension of the “enhanced production zone” is drastically reduced and even bacteria could not benefit from the fast mixing regime induced.     

Description of a flexible and extendable physical–biogeochemical model system for the water column

A modelling system for coupled physical–biogeochemical simulations in the water column is presented here. The physical model component allows for a number of different statistical turbulence closure schemes, ranging from simple algebraic closures to two-equation turbulence models with algebraic second-moment closures. The biogeochemical module consists of models which are based on a number of state variables represented by their ensemble averaged concentrations. Specific biogeochemical models may range from simple NPZ (nutrient–phytoplankton–zooplankton) to complex ecosystem models. Recently developed modified Patankar solvers for ordinary differential equations allow for stable discretisations of the production and destruction terms guaranteeing conservative and non-negative solutions. The increased stability of these new solvers over explicit solvers is demonstrated for a plankton spring bloom simulation. The model system is applied to marine ecosystem dynamics the Northern North Sea and the Central Gotland Sea. Two different biogeochemical models are applied, a conservative nitrogen-based model to the North Sea, and a more complex model including an oxygen equation to the Baltic Sea, allowing for the reproduction of chemical processes under anoxic conditions. For both applications, earlier model results obtained with slightly different model setups could be basically reproduced. It became however clear that the choice for ecosystem model parameters such as maximum phytoplankton growth rates does strongly depend on the physical model parameters (such as turbulence closure models or external forcing).     

海洋生态系统及海岸工程生态影响预测模型研究进展

对海洋生态系统的研究已形成了以实验观测为基础、建立生态模型的方法。文章对已有的海洋生态模型研究成果进行了归纳总结,认为初级生产力模型是纯生态模型的主要发展方向,综合考虑物理生化作用的生态系统动力学模型将是未来海洋生态研究的必然趋势。同时,由于我国大规模海岸工程建设对近岸海域生态系统造成了较大影响,而相关的生态影响预测模型研究较少,为有效预测评价海岸工程建设对近岸海域生态系统的影响,进一步深入研究海岸工程建设对近岸海洋生态影响预测模型将显得尤为重要。     

Modelling Pseudocalanus elongatus stage-structured population dynamics embedded in a water column ecosystem model for the northern North Sea

This paper outlines an approach to couple a structured zooplankton population model with state variables for eggs, nauplii, two copepodites stages and adults adapted to Pseudocalanus elongatus into the complex marine ecosystem model ECOHAM2 with 13 state variables resolving the carbon and nitrogen cycle. Different temperature and food scenarios derived from laboratory culture studies were examined to improve the process parameterisation for copepod stage dependent development processes. To study annual cycles under realistic weather and hydrographic conditions, the coupled ecosystem–zooplankton model is applied to a water column in the northern North Sea. The main ecosystem state variables were validated against observed monthly mean values. Then vertical profiles of selected state variables were compared to the physical forcing to study differences between zooplankton as one biomass state variable or partitioned into five population state variables. Simulated generation times are more affected by temperature than food conditions except during the spring phytoplankton bloom. Up to six generations within the annual cycle can be discerned in the simulation.     

Environmental changes in the western Black Sea related to anthropogenic and natural conditions

The Black Sea northwestern shelf (NWS) ecosystem has been subjected to the strongest anthropogenic pressure of the entire Black Sea as about 80% of the freshwater runoff is discharged there.This paper presents a review of the global environmental changes related mainly to increased eutrophication in the western Black Sea basin.A case study (CoMSBlack 92a cruise) attempts to highlight the interaction of some natural and anthropogenic factors responsible for specific chemical and biological features in the western Black Sea environment. The significance of processes located near the Danube river mouth, which, due to their relatively small space scale, have not been recognized before, is clarified. Hydrological processess of specific importance to the distribution and transformation of antropogenic inputs are river plume dynamics, coastal upwelling and mixing and downwelling over the shelf break and slope in this part of the basin. A layer of low hydrological variability (“conservative layer”) appears to be a natural feature of the area, hypothesized to precondition specific biological and chemical processes there.The results suggest that the interplay between the Danube anthropogenic nutrient load with the natural hydrological fronts and gradients provides opportunities for enhanced biological activity thus contributing to the global environmental changes in the Black Sea NWS.     

Biological effects of dilution of Dead Sea brine with seawater: implications for the planning of the Red Sea–Dead Sea “Peace Conduit”

The Dead Sea is a severely disturbed ecosystem. Its water level has been decreasing at a rate of nearly 1 m per year during the last decade due to anthropogenic intervention in its water balance. Since the peace treaty between Israel and Jordan was established in 1994, a proposal for the construction of a water carrier, the “Peace Conduit,” between the Gulf of Aqaba (Red Sea) and the Dead Sea is being investigated. This water carrier is intended to mitigate damaging processes that currently occur in the Dead Sea and its surrounding area. The difference in elevation of about 416 m could be exploited for seawater desalination by reverse osmosis. To examine the possible effects of the mixing of Dead Sea brines with seawater and seawater concentrates on the Dead Sea as an ecosystem, we have set up simulation experiments under field conditions in experimental ponds at Sedom, in which Dead Sea water was diluted with Red Sea water. The main components of the Dead Sea biota are the unicellular green alga Dunaliella and several types of red halophilic Archaea. Phosphate is the limiting inorganic nutrient. Massive Dunaliella blooms developed, accompanied by dense communities of red halophilic Archaea, in some of the experimental ponds, imparting a brown-red coloration to the brines. The extent of biological development depended on the extent of dilution and on phosphate availability. The results of the simulation experiments show that biological phenomena and their impact on the Dead Sea ecosystem should be taken into consideration during the planning of the “Peace Conduit.”     

An eddy-permitting, coupled ecosystem-circulation model of the Arabian Sea: comparison with observations

A nitrogen-based, pelagic ecosystem model has been coupled with an eddy-permitting ocean general circulation model of the Arabian Sea, and the results are compared with observations. The seasonal variability simulated by the model is in good agreement with observations: during the southwest monsoon season, phytoplankton increases in the western Arabian Sea due to upwelling along the coast; during the northeast monsoon season, phytoplankton abundance is large in the northern Arabian Sea because of the enhanced nitrate entrained by relatively deep vertical mixing. Two major differences are, however, found in the basin-wide comparison between model results and observations: an unrealistic nitrate maximum in the subsurface layer of the northern Arabian Sea and too low primary production in oligotrophic regimes. The former may be attributed to the lack of denitrification in the model. Possible causes for the latter include the present model's underestimation of fast nutrient recycling, the neglect of carbon fixation decoupled from nitrogen uptake and of nitrogen fixation, and inadequate nitrate entrainment by mixed layer deepening. The rate at which simulated nitrate increases in the northern Arabian Sea is 11–24 TgN/year, and should correspond to the denitrification rate integrated over the northern Arabian Sea assuming that the loss of nitrogen through denitrification is balanced by advective input. The model does not reproduce the observed phytoplankton bloom in the late southwest monsoon season. Possible causes are that the mixed layer may be too shallow in summer and that the horizontal transport of nitrate from the coast of Oman may be too weak. Sensitivity experiments demonstrate a strong dependence of the simulated primary productivity on the vertical mixing scheme and on the inclusion of a fast recycling loop in the ecosystem model.     

Weaving marine food webs from end to end under global change

Marine food web dynamics are determined by interactions within and between species and between species and their environment. Global change directly affects abiotic conditions and living organisms, impinging on all trophic levels in food webs. Different groups of marine researchers traditionally study different aspects of these changes. However, over medium to long time scales perturbations affecting food webs need to be considered across the full range from nutrients to top predators. Studies of end-to-end marine food webs not only span organism sizes and trophic levels, but should also help align multidisciplinary research to common goals and perspectives. Topics are described that bridge disciplinary gaps and are needed to develop new understanding of the reciprocal impacts of global change on marine food webs and ocean biogeochemistry. These include (1) the effects of nutrients on biomass and production, (2) the effects of varying element ratios on food web structure and food quality, (3) bulk flows of energy and material in food webs and their efficiencies of transfer, (4) the ecological effects of species richness and the roles of microbial organisms, (5) the role of feeding behaviour in food web dynamics and trophic controls, (6) the spatial dynamics of communities and links between different food webs, (7) the combined effects of body size and behaviour in determining dynamics of food webs, and (8) the extent to which the ability of marine organisms (and communities) to adapt will influence food web dynamics. An overriding issue that influences all topics concerns the time and space scales of ecosystem variability. Threads link different nodes of information among various topics, emphasizing the importance of tackling food web studies with a variety of modelling approaches and through a combination of field and experimental studies with a strong comparative approach.     

The small-sized benthic biota of the Håkon Mosby Mud Volcano (SW Barents Sea slope)

R/V POLARSTERN expedition ARK XVIII/1 in summer 2002 provided the opportunity to carry out a sampling programme to assess the activity, biomass and composition of the small-sized benthic biota (size range: bacteria to meiofauna) around the active mud-oozing and methane-seeping Håkon Mosby Mud Volcano (HMMV) on the SW Barents Sea slope, Northern North-Atlantic. A total of 11 stations, covering different areas (e.g., bacterial mat sites, and pogonophoran fields) within the crater, and sites outside the caldera were sampled using a multiple corer. Subsamples were analyzed for various biogenic compounds to estimate the flux of organic matter to the seafloor (sediment-bound chloroplastic pigments indicating phytodetritus), activities (bacterial exo-enzymatic turnover rates) and the total biomass [from bulk parameters, like phospholipid (PL) concentrations in the sediments] of the smallest sediment-inhabiting organisms (range: bacteria to meiofauna). Direct investigations of bacterial numbers and biomasses as well as on meiofauna densities and composition completed our investigations at HMMV. As expected for a comparable small deep-sea area with only minor disparity in water depth between sampling sites, our investigations revealed generally no significant differences in organic matter input from phytodetritus sedimentation between sampling sites inside and outside HMMV. Bacterial exo-enzymatic activities as well as total microbial biomass (TMB) and meiofauna densities, however, exhibited generally higher values at HMMV, compared to sites outside the mud volcano. Enhanced benthic life at HMMV is based on chemosynthetic processes, making the mud volcano a “chemosynthetic oasis” in an otherwise oligotrophic deep-sea environment. As we did not find any indication for bacterial symbioses in the meiofauna, comparably rich meiofaunal assemblages at HMMV are presumably indirectly related to a general enhanced biological production.     

Scientific requirements for an abyssal benthic laboratory

For over thirty years man has studied “outer space” and installed satellites which watch the surface of the Earth. The great depths of the world ocean are, however, practically unknown and there is an urgent need to put abyssal benthic laboratories into “inner space” in order to study basic phenomena of interest to marine science and climatology as well as man's impact on the oceans.In view of the numerous problems related to global change, as a first step emphasis should first be on the role of the oceans and their inherent processes, which are the focus of such international programmes as the World Ocean Circulation Experiment (WOCE) and the Joint Global Ocean Flux Study (JGOFS). Multi-disciplinary registration of key events at selected key sites investigating the variability in time and space are of the utmost importance. The same methods and techniques must be used for the study of human impacts on the deep oceans caused by mining of metalliferous resources and by waste disposal as well as in basic studies. However, the investigation of the inner space of our planet has certain requirements. As long-term and large-scale investigations become more and more important, development of automized systems, largely independent from research vessels will be required. This will demand high capacities of energy for all technical functions as well as high storage capacities for data and samples. As a consequence the needs for two different—although overlapping—functional approaches are defined for future deep-sea deployments.(A) A system for long-term registration of the natural variability and long-term monitoring of human impacts: (B) A system for short-term observations and short-time experimentations. This report summarizes their technological demands. The envisioned interdisciplinary technology should deliver information on physical, biological and geochemical processes and their variabilities in the deep oceans. The prospected systems need to have the ability for real time video observation, data transfer and experimental manipulation, as well as sensing and sampling facilities with large storage capacities for long-term deployments.Prospective costs of the described multipurpose abyssal benthic laboratory will presumably exceed the funds for deep-sea research of a single country. A joint European effort could solve this problem and help to manifest a leading role for European marine science in international deep-sea and global change research.     

The importance of polynyas, ice edges, and leads to marine mammals and birds

The correlation between areas of open water in ice-covered seas and increased biological productivity has been noted for some time. To date, most attention has been focused on larger polynyas, such as the Northeast Water and the Northwater. Although spectacular in their own right, these large polynyas represent only part of a vitally important continuum of biological productivity that varies significantly between geographic areas and ice habitats, that includes the multi-year pack of the polar ocean and small localized polynyas in annual ice. Surveys of the distribution and abundance of ringed seals in the Canadian Arctic Archipelago have shown differences in density that are correlated with the presence or absence of polynyas. There is also significant variation in the biological productivity of polynya areas of the Canadian High Arctic Archipelago and northern Greenland, all of which receive inflow from the polar basin. Long-term studies of polar bears and ringed seals in western Hudson Bay and the eastern Beaufort Sea show significant but dissimilar patterns of change in condition and reproductive rates between the two regions and suggest that fundamentally different climatic or oceanographic processes may be involved. Projections of climate models suggest that, if warming occurs, then the extent of ice cover in Hudson Bay may be among the first things affected. Long-term studies of polar bears and ringed seals in the eastern Beaufort Sea and Hudson Bay would suggest these two species to be suitable indicators of significant climatic or oceanographic changes in the marine ecosystem.     

Seasonal and interannual variation of physical and biological processes during 1994–2001 in the Sea of Japan/East Sea: A three-dimensional physical–biogeochemical modeling study

A Pacific basin-wide physical–biogeochemical model has been used to investigate the seasonal and interannual variation of physical and biological fields with analyses focusing on the Sea of Japan/East Sea (JES). The physical model is based on the Regional Ocean Model System (ROMS), and the biogeochemical model is based on the Carbon, Si(OH)₄, Nitrogen Ecosystem (CoSiNE) model. The coupled ROMS–CoSiNE model is forced with the daily air–sea fluxes derived from the National Centers for Environmental Prediction (NCEP) and the National Center for Atmospheric Research (NCAR) reanalysis for the period of 1994 to 2001, and the model results are used to evaluate climate impact on nutrient transport in Mixed Layer Depth (MLD) and phytoplankton spring bloom dynamics in the JES.The model reproduces several key features of sea surface temperature (SST) and surface currents, which are consistent with the previous modeling and observational results in the JES. The calculated volume transports through the three major straits show that the Korea Strait (KS) dominates the inflow to the JES with 2.46 Sv annually, and the Tsugaru Strait (TS) and the Soya Strait (SS) are major outflows with 1.85 Sv and 0.64 Sv, respectively. Domain-averaged phytoplankton biomass in the JES reaches its spring peak 1.8 mmol N m^− 3 in May and shows a relatively weak autumn increase in November. Strong summer stratification and intense consumption of nitrate by phytoplankton during the spring result in very low nitrate concentration at the upper layer, which limits phytoplankton growth in the JES during the summer. On the other hand, the higher grazer abundance likely contributes to the strong suppression of phytoplankton biomass after the spring bloom in the JES. The model results show strong interannual variability of SST, nutrients, and phytoplankton biomass with sudden changes in 1998, which correspond to large-scale changes of the Pacific Decadal Oscillation (PDO). Regional comparisons of interannual variations in springtime were made for the southern and northern JES. Variations of nutrients and phytoplankton biomass related to the PDO warm/cold phase changes were detected in both the southern and northern JES, and there were regional differences with respect to the mechanisms and timing. During the warm PDO, the nutrients integrated in the MLD increased in the south and decreased in the north in winter. Conversely, during the cold PDO, the nutrients integrated in the MLD decreased in the south and increased in the north. Wind divergence/convergence likely drives the differences in the southern and northern regions when northerly and northwesterly monsoon dominates in winter in the JES. Subjected to the nutrient change, the growth of phytoplankton biomass appears to be limited neither by nutrient nor by light consistently both in the southern and northern regions. Namely, the JES is at the transition zone of the lower trophic-level ecosystem between light-limited and nutrient-limited zones.     

A generalized model of pelagic biogeochemistry for the global ocean ecosystem. Part II: Numerical simulations

This paper presents a global ocean implementation of a multi-component model of marine pelagic biogeochemistry coupled on-line with an ocean general circulation model forced with climatological surface fields (PELAgic biogeochemistry for Global Ocean Simulations, PELAGOS). The final objective is the inclusion of this model as a component in an Earth System model for climate studies. The pelagic model is based on a functional stoichiometric representation of marine biogeochemical cycles and allows simulating the dynamics of C, N, P, Si, O and Fe taking into account the variation of their elemental ratios in the functional groups. The model also includes a parameterization of variable chlorophyll/carbon ratio in phytoplankton, carrying chl as a prognostic variable. The first part of the paper analyzes the contribution of non-local advective–diffusive terms and local vertical processes to the simulated chl distributions. The comparison of the three experiments shows that the mean chl distribution at higher latitudes is largely determined by mixing processes, while vertical advection controls the distribution in the equatorial upwelling regions. Horizontal advective and diffusive processes are necessary mechanisms for the shape of chl distribution in the sub-tropical Pacific. In the second part, the results have been compared with existing datasets of satellite-derived chlorophyll, surface nutrients, estimates of phytoplankton community composition and primary production data. The agreement is reasonable both in terms of the spatial distribution of annual means and of the seasonal variability in different dynamical oceanographic regions. Results indicate that some of the model biases in chl and surface nutrients distributions can be related to deficiencies in the simulation of physical processes such as advection and mixing. Other discrepancies are attributed to inadequate parameterizations of phytoplankton functional groups. The model has skill in reproducing the overall distribution of large and small phytoplankton but tends to underestimate diatoms in the northern higher latitudes and overestimate nanophytoplankton with respect to picoautotrophs in oligotrophic regions. The performance of the model is discussed in the context of its use in climate studies and an approach for improving the parameterization of functional groups in deterministic models is outlined.     

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.