Simulation of coupled pelagic-benthic ecosystem of the Yellow Sea Cold Water Mass

A one-dimensional coupled pelagic-benthic box model for the Yellow Sea Cold Water Mass （YSCWM） is developed. The model is divided into three boxes vertically according to the depths of thermocline and euphotic layer. It simulates well the oligotrophic shelf ecosystem of the YSCWM considering effects of nutrients deposition and microbial loop. Main features of vertical structure of various variables in ecosystem of the YSCWM were captured and seasonal variability of the ecosystem was well reconstructed. Calculation shows that the contribution of microbial loop to the zooplankton can reach up to 60%. Besides, input of inorganic nutrients from atmospheric deposition is an important mechanism of production in upper layer of the YSCWM when stratified.     

COUPLED PHYSICAL-ECOLOGICAL MODELLING IN THE CENTRAL PART OF JIAOZHOU BAY Ⅱ.COUPLED WITH AN ECOLOGICAL MODEL

Sharpies‘ 1-D physical rrozlel maploying tide-wind driven turbulence closure and surface heating-cooling physics, was coupled with an eculogical rnodet with 9-biochemical components: phytoplankton, zooplankton, shellfish, autotmphic and heterotrophic bacterioplankton, dissolved organic carbon (DOC), suspended detritus and sinking particles to simulate the armual evolution of ecosystem in thecentral part of Jiaozhou Bay. The coupled modeling results showed that the phytoplankton shading effectcould reduce seawater temperamre by 2℃, so that photosynthesis efficiency should be less than 8% ; that the loss of phytoplankton by zooplankton grazing in winter tended to be compensated by phytoplankton advection and diffusion from the otrtside of the Bay; that the incidem irradiance intensity could be the mostimportant factor for phytoplankton grcr, wth rate; and that it was the bacterial secondary prnduction that maintained the maximum zooplankton biomass in winter usually observed in the 1990s, indicating that themicrobial food loop was extremely important for ecosystem study of Jiaozhou Bay.     

Simple ecosystem model of the central part of the East China Sea in spring

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Pelagic Nitrogen Cycling in Jiaozhou Bay, a Model Study I： the Conceptual Model

A zero-dimensional box model (PNCMjzb) with six state variables (ammonium, nitrate, dissolved organic nitrogen, phytoplankton, zooplankton and detritus) was developed to study nitrogen cycling in the Jiaozhou Bay pelagic ecosystem. The dominant processes within these compartments are considered with nitrogen as flow currency. Phytoplankton and zooplankton are treated as separate state variables, assuming that the species composition was dominated by two or three species the dynamic constants of which are similar and that they represent the entire plankton community. The microbial loop has not been integrated explicitly in the model. The turnover of bacteria is included implicitly in processes such as detritus decomposition, DON remineralization, pelagic nitrification and denitrification. The model is driven by two forcing variables, viz. water temperature and light intensity. Historical data from the1980s and 1990s were compiled and used for model calibration. In this paper (part Ⅰ), the consideration of every main compartment in the model is interpreted in detail. And the applied equations and parameters are presented. The main results from the simulations together with discussion about phytoplankton dynamics and primary production in Jiaozhou Bay are presented in the next paper (part Ⅱ).     

Diagnosis of physical and biological controls on phytoplankton distribution in the Sargasso Sea

The linkage between physical and biological processes is studied by applying a one-dimensional physical-biological coupled model to the Sargasso Sea. The physical model is the Princeton Ocean Model and the biological model is a five-component system including phytoplankton, zooplankton, nitrate, ammonium, and detritus. The coupling between the physical and biological model is accomplished through vertical mixing which is parameterized by the level 2.5 Mellor and Yamada turbulence closure scheme. The coupled model investigates the annual cycle of ecosystem production and the response to external forcing, such as heat flux, wind stress, and surface salinity, and the relative importance of physical processes in affecting the ecosystem. Sensitivity experiments are also carried out, which provide information on how the model bio-chemical parameters affect the biological system. The computed seasonal cycles compare reasonably well with the observations of the Bermuda Atlantic Time-series Study (BATS). The spring bloom of phytoplankton occurs in March and April, right after the weakening of the winter mixing and before the establishment of the summer stratification. The bloom of zooplankton occurs about two weeks after the bloom of phytoplankton. The sensitivity experiments show that zooplankton is more sensitive to the variations of biochemical parameters than phytoplankton.     

A coupled ice-ocean ecosystem model for 1-D and 3-D applications in the Bering and Chukchi Seas

Primary production in the Bering and Chukchi Seas is strongly influenced by the annual cycle of sea ice. Here pelagic and sea ice algal ecosystems coexist and interact with each other. Ecosystem modeling of sea ice associated phytoplankton blooms has been understudied compared to open water ecosystem model applications. This study introduces a general coupled ice-ocean ecosystem model with equations and parameters for 1-D and 3-D applications that is based on 1-D coupled ice-ocean ecosystem model development in the landfast ice in the Chukchi Sea and marginal ice zone of Bering Sea. The biological model includes both pelagic and sea ice algal habitats with 10 compartments： three phytoplankton （pelagic diatom, flagellates and ice algae： D, F, and Ai） , three zooplankton （copepods, large zooplankton, and microzooplankton ： ZS, ZL, ZP） , three nutrients （ nitrate ＋ nitrite, ammonium, silicon ： NO3 , NH4, Si） and detritus （Det）. The coupling of the biological models with physical ocean models is straightforward with just the addition of the advection and diffusion terms to the ecosystem model. The coupling with a multi-category sea ice model requires the same calculation of the sea ice ecosystem model in each ice thickness category and the redistribution between categories caused by both dynamic and thermodynamic forcing as in the physical model. Phytoplankton and ice algal self-shading effect is the sole feedback from the ecosystem model to the physical model.     

Trophic interactions, ecosystem structure and function in the southern Yellow Sea

The southern Yellow Sea is an important fishing ground, providing abundant fishery resources. However, overfishing and climate change have caused a decline in the resource and damaged the ecosystem. We developed an ecosystem model to analyze the trophic interactions and ecosystem structure and function to guide sustainable development of the ecosystem. A trophic mass-balance model of the southern Yellow Sea during 2000–2001 was constructed using Ecopath with Ecosim software. We defined 22 important functional groups and studied their diet composition. The trophic levels of fish, shrimp, crabs, and cephalopods were between 2.78 and 4.39, and the mean trophic level of the fisheries was 3.24. The trophic flows within the food web occurred primarily in the lower trophic levels. The mean trophic transfer efficiency was 8.1%, of which 7.1% was from primary producers and 9.3% was from detritus within the ecosystem. The transfer efficiency between trophic levels II to III to IV to V to >V was 5.0%, 5.7%, 18.5%, and 19.7%–20.4%, respectively. Of the total flow, phytoplankton contributed 61% and detritus contributed 39%. Fishing is defined as a top predator within the ecosystem, and has a negative impact on most commercial species. Moreover, the ecosystem had a high gross efficiency of the fishery and a high value of primary production required to sustain the fishery. Together, our data suggest there is high fishing pressure in the southern Yellow Sea. Based on analysis of Odum’s ecological parameters, this ecosystem was at an immature stage. Our results provide some insights into the structure and development of this ecosystem.     

A model study on dynamical processes of phytoplankton in Laizhou Bay

A three-dimensional ecosystem model, using a PIC (Particle-In-Cell) method, is developed to reproduce the annual cycle and seasonal variation of nutrients and phytoplankton biomass in Laizhou Bay. Eight state variables, i.e., DIN (dissolved inorganic nitrogen), phosphate, DON (dissolved organic nitrogen), DOP (dissolved organic phosphorus), COD (chemical oxygen demand), chlorophyll-a (Chl-a), detritus and the zooplankton biomass, are included in the model. The model successfully reproduces the observed temporal and spatial variations of nutrients and Chl-a biomass distributions in the bay. The nutrient concentrations are at high level in winter and at low level in summer. Double-peak structure of the phytoplankton (PPT) biomass exists in Laizhou Bay, corresponding to a spring and an autumn bloom respectively. Several numerical experiments are carried out to examine the nutrient limitation, and the importance of the discharges of the Yellow River and Xiaoqinghe River. Both DIN limitation and phosphate limitation exist in some areas of the bay, with the former being more significant than the latter. The Yellow River and Xiaoqinghe River are the main pollution sources of nutrients in Laizhou Bay. During the flood season, the algal growth is inhibited in the bay with the Yellow River discharges being excluded in the experiment, while in spring, the algal growth is enhanced with the Xiaoqinghe River excluded.     

Adjoint Assimilation in Marine and an Example of Application

This paper aims at a review of the work carried out to date on the adjoint assimilation of data in marine ecosystem models since 1995. The structure and feature of the adjoint assimilation in marine ecosystem models are also introduced. To illustrate the application of the adjoint technique and its merits, a 4-variable ecosystem model coupled with a 3-D physical model is established for the Bohai Sea and the Yellow Sea. The chlorophyll concentration data derived from the SeaWiFS ocean colour data are assimilated in the model with the technique. Some results are briefly presented.     

Benthic nutrient recycling in shallow coastal waters of the Bohai Sea

Sediment-water fluxes of N and P species in the Bohai Sea were investigated in September-October 1998 and April-May 1999. The benthic fluxes of nutrient species were determined by incubating sediment core samples with bottom seawater bubbled with air or nitrogen. NO^-2,NH4, dissolved organic nitrogen (DON) and phosphorus (DOP), total dissolved nitrogen (TDN) and phosphorus (TDP), and PO4^3- showed a net exchange flux from seawater to sediment, while NO^-3, dissolved inorganic nitrogen (DIN) and SiO3^2- were released from sediment to seawater in the Bohai Sea. Sediment-water nutrient exchange increases DIN and reduces the phosphorus load in the Bohai Sea. The release of silicate from sediment to overlying seawater reduces potential silicate limitation of primary production resulted from decrease of riverine discharge.The exchange flux of nutrients showed no obvious seasonal variation. The present study showed that the concentrations and composition of nutrients in the water column were affected by suspended sediment, and that not all the exchangeable phosphate in sediment could be released via sediment resuspension.     

Estimates of autumntime benthic secondary production in Laizhou Bay and adjacent Bohai Sea waters

The whole metazoan community inhabiting Laizhou Bay and adjacent Bohai Sea waters were sampled in late autumn,2006.Secondary production estimates for macrofauna and meiofauna were made separately.Total benthic secondary production was as high as 8.38 ± 4.08 g ash-free dry weight(AFDW) m-2 a-1,which represented the autumn production level.In general,macrofaunal secondary production in Laizhou Bay was much lower than that in adjacent Bohai Sea areas.In contrast,meiofaunal secondary production in Laizhou Bay was higher than that in adjacent Bohai Sea areas.Macrofauna contributed 61% to benthic secondary production(5.09 ± 3.26 g AFDW m-2 a-1),lower than the value in previous studies in Bohai Sea.Sediment granulometric characteristics and bottom-water salinity could explain the substantial variability in the macrofauna biomass and production.Meiofaunal production was an important component of benthic production and exceeded macrofauna production under exceptional conditions,e.g.in Laizhou Bay,where macrofauna was restricted.Chlorophyll pigments(Chl-a) concentrations in sediment explained the general meiofaunal biomass and production distribution here.     

Changes in nitrogen and phosphorus and their effects on phytoplankton in the Bohai Sea

Systematic studies of the changes in dissolved inorganic nitrogen (DIN) and dissolved inorganic phosphorus (DIP) and their effects on phytoplankton over the last 30 years in the Bohai Sea are presented. The amount of sewage disposal, use of fertilizer and the Huanghe River runoff were found to have a significant influence on the DIN or DIP concentrations in the Bohai Sea over the last 30 years. Moreover, the changes in DIN and DIP resulted in changes in the limiting nutrients of phytoplankton in the Bohai Sea from nitrogen in the early 1980s to nitrogen-phosphorus in the late 1980s, and then to phosphorus after the 1990s. In addition, changes in nitrogen and phosphorus had a significant effect on the phytoplankton community structure. The half saturation constant (K _s) was used to evaluate the effect of nutrients on the phytoplankton community structure in the Bohai Sea over the last 30 years. Cell abundance percentages of dominant phytoplankton species with high K _s values for phosphorus and low K _s values for nitrogen have decreased since the 1980s, while those of dominant phytoplankton species with low K _s values for phosphorus and high K _s values for nitrogen increased during this period.     

Taxonomic distinctness of macrofauna as an ecological indicator in Laizhou Bay and adjacent waters

In this paper, we assessed the ecological and biodiversity status in the Bohai Sea through a quantitative survey on mac-rofaunal community at 25 stations in Laizhou Bay and adjacent waters in the autumn of 2006.We tested the robustness and effectiveness of taxonomic distinctness as an ecological indictor by analyzing its correlation with species richness and natural environmental variables and by analyzing other ecological indicators (Shannon-Wiener H' and W statistics from Abundance Biomass Comparison curve).Results so obtained indicated that the benthic environment of the study waters in general is not under major impact of anthropogenic disturbance, but some stations in Laizhou Bay and along the coast of the Shandong Peninsula and even in the central Bohai Sea might be moderately disturbed and showed signs of ecological degradation.The taxonomic distinctness measures △+ and Λ+ were independent of sampling effort and natural environment factors and were compliant to other ecological indicators.Further application of the taxonomic distinctness indicator to assess marine biodiversity and ecosystem health on a larger regional scale with historical data seems promising.     

Seasonal dynamics of crustacean zooplankton community structure in Erhai Lake,a plateau lake,with reference to phytoplankton and environmental factors

The seasonal dynamics of a crustacean zooplankton community in Erhai Lake was investigated from May 2010 to April 2011. In total, 11 species were recorded, including six （6 genera） cladoceran and five （5 genera） copepod species. The crustacean zooplankton densities ranged from 24.3 to 155.4 ind./L. In winter and spring, the large-bodied cladoceran Daphnia galeata dominated the crustacean plankton community. In summer and autumn, when the colonial or filamentous algae dominated the phytoplankton communities, the small-bodied species （e.g. Bosminafatalis, Ceriodaphnia quadrangular, and Mesocyclops leuckarti） replaced the large-bodied ones. One-way ANOVA and redundancy analysis revealed that community structure was dependent upon total nitrogen, total phosphorus, water temperature, transparency, and the biomass of small algae. The variation in both phytoplankton structure and environmental variables were important factors in the seasonal succession of crustacean zooplankton structure in Erhai Lake.     

Diel vertical migration of mesozooplankton in the northern Yellow Sea

The Yellow Sea Cold Water Mass(YSCWM),one of the most vital hydrological features of the Yellow Sea,causes a seasonal thermocline from spring to autumn.The diel vertical migration(DVM) of zooplankton is crucial to structural pelagic communities and food webs,and its patterns can be affected by thermocline depth and strength.Hence,we investigated zooplankton community succession and seasonal changes in zooplankton DVM at a fixed station in the YSCWM.Annual zooplankton community succession was affected by the forming and fading of the YSCWM.A total of 37 mesozooplankton taxa were recorded.The highest and lowest species numbers in autumn and spring were detected.The highest and lowest total densities were observed in autumn(14 464.1 inds./m3) and winter(3 115.4 inds./m3),respectively.The DVM of the dominant species showed obvious seasonal variations.When the YSCWM was weak in spring and autumn,most species(e.g.Paracalanus parvus,Oithona similis,and Acartia bifilosa) stayed above the thermocline and vertically migrated into the upper layer.Calanus sinicus and Aidanosagitta crassa crossed the thermocline and vertically migrated.No species migrated through the stratification in summer,and all of the species were limited above(P.parvus and A.crassa) or below(C.sinicus and Centropages abdominalis)the thermocline.The YSCWM disappeared in winter,and zooplankton species were found throughout the water column.Thus,the existence of thermocline influenced the migration patterns of zooplankton.Cluster analyses showed that the existence of YSCWM resulted in significant differences between zooplankton communities above and below the thermocline.     

Seasonal Variation of Environmental Variables and Phytoplankton Community Structure and Their Relationship in Liaodong Bay of Bohai Sea,China

In this paper, systematic studies on the changes in concentrations of the environmental factors and the net-phytoplankton community, and the relationship between them in the Liaodong Bay, Bohai Sea during 2013 are presented. The PCA results showed that higher levels of nutrients and dissolved heavy metals in the river-estuary-bay system were closely related to the river runoff. Since the influences of industrial and anthropogenic activities, the Liaodong Bay coastal areas are facing a huge environmental challenge of nutrients and heavy metal pollution. Net-phytoplankton community structure showed obvious seasonal succession, among which the dominant and (or) key species were the main factors affecting community structure change and stability. Under certain environmental conditions, the dominant species and (or) key species dominated the phytoplankton community structure succession. The Bio-ENV results suggested that the seawater temperature, nutrient, Cu, Pb, Zn, and Cd in Liaodong Bay are important environmental variables that affect the phytoplankton community structure. Anthropogenic activities have significantly contributed to the changes in concentrations of environmental factors and the net-phytoplankton community structure and stability, and the relationship between them.     

Coupling of an individual-based model of anchovy with lower trophic level and hydrodynamic models

Anchovy (Engraulis japonicus), a small pelagic fish and food of other economic fishes, is a key species in the Yellow Sea ecosystem. Understanding the mechanisms of its recruitment and biomass variation is important for the prediction and management of fishery resources. Coupled with a hydrodynamic model (POM) and a lower trophic level ecosystem model (NEMURO), an individual-based model of anchovy is developed to study the influence of physical environment on anchovy’s biomass variation. Seasonal variations of circulation, water temperature and mix-layer depth from POM are used as external forcing for NEMURO and the anchovy model. Biomasses of large zooplankton and predatory zooplankton which anchovy feeds on are output from NEMURO and are controlled by the consumption of anchovy on them. Survival fitness theory related to temperature and food is used to determine the swimming action of anchovy in the model. The simulation results agree well with observations and elucidate the influence of temperature in over-wintering migration and food in feeding migration.     

DISTRIBUTION AND POPULATION DYNAMICS OF PARACALANUS PARVUS, PARACALANUS CRASSIROSTRIS, AND ACARTIA BIFILOSA (COPEPODA, CALANOIDA) IN THE BOHAI SEA

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The community structure and seasonal dynamics of plankton in Bange Lake,northern Tibet,China

The seasonal variations in biomass, abundance, and species composition of plankton in relation to hydrography were studied in the saline Bange Lake, northern Tibet, China. Sampling was carried out between one to three times per month from May 2001 to July 2002. Salinity ranged from 14 to 146. The air and water temperature exhibited a clear seasonal pattern, and mean annual temperatures were approximately 4.8°C and 7.3°C, respectively. The lowest water temperature occurred in winter from December to March at-2°C and the highest in June and July at 17.7°C. Forty-one phytoplankton taxa, 21 zooplankton, and 5 benthic or facultative zooplankton were identifi ed. The predominant phytoplankton species were Gloeothece linearis, Oscillatoria tenuis, Gloeocapsa punctata, Ctenocladus circinnatus, Dunaliella salina, and Spirulina major. The predominant zooplankton species included H olophrya actra, Brachionus plicatilis, Daphniopsis tibetana, Cletocamptus dertersi, and A rctodiaptomus salinus. The mean annual total phytoplankton density and biomass for the entire lake were 4.52×10~7 cells/L and 1.60 mg/L, respectively. The annual mean zooplankton abundance was 52, 162, 322, and 57, 144 ind./L, in the three sublakes. The annual mean total zooplankton biomass in Lakes 1–3 was 1.23, 9.98, and 2.13 mg/L, respectively. The annual mean tychoplankton abundances in Bg1, 2, and 3 were 47, 67, and 654 ind./L. The annual mean tychoplankton biomass was 2.36, 0.16, and 2.03 mg/L, respectively. The zooplankton biomass(including tychoplankton) in the lake was 9.11 mg/L. The total number of plankton species in the salt lake was signifi cantly negatively correlated with salinity.     

Modelling nitrogen and phosphorus dynamics in a mesocosm pelagic ecosystem in Laizhou Bay in China

A model of nitrogen and phosphorus dynamics in mesocosm experiments was established on the basis of a summary and synthesis of the existing models. The established model comprised seven state variables(DIN,PO4-P,DON,DOP,phytoplankton,zooplankton and detritus) and five modules - phytoplankton,zooplankton,dissolved inorganic nutrients,dissolved organic nutrients and detritus. Comparison with the in situ experimental data in Laizhou Bay at the end of August 2002 showed that this model could properly simulate t...