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Obvious spatiotemporal heterogeneity is a distinct characteristic of ecosystems in subtropical bays. To aid targeted management and ecological restoration in long and narrow semi-enclosed subtropical bays, we analyzed seasonal and regional differences in long-term changes (1980–2019) in the biomass and abundance of large mesozooplankton (LMZ; >505 µm) in Xiangshan Bay, Zhejiang, China. We found spatiotemporal heterogeneity in the historical changes of LMZ. Significant negative trends in LMZ biomass were found in the inner and middle bay during the warm season (summer and autumn), when the nutrient concentration (especially dissolved inorganic nitrogen) and temperature increased simultaneously. Nutrient changes in Xiangshan Bay began in the late 1980s or early 1990s, coinciding with large-scale fish cage development. A rapid decline in LMZ biomass occurred after 2005 when power plants commenced operation, accelerating the warming trend. Therefore, the joint stress of eutrophication and warming likely precipitated the decline in LMZ biomass. Conversely, a significant increase in LMZ biomass was found in the outer bay in spring. This trend was consistent with the trend of LMZ biomass near the Changjiang (Yangtze) River estuary, which indicates that the pelagic ecosystem in the outer bay was affected by water from the Changjiang River estuary during spring. Based on our results, ecosystem management and restoration in semi-enclosed subtropical bays should focus on internal waters, which have a poor capacity for water exchange. For Xiangshan Bay, the changes in the Changjiang River estuary ecosystem during the cold season (winter and spring) should also be considered.

Temperature increase caused by thermal discharge from power plants promotes phytoplankton growth and frequent bloom in eutrophic subtropical waters, particularly in cold seasons. Suspension filter-feeding bivalves show size-selective grazing on phytoplankton. Thus, we hypothesized that algal bloom under thermal stimulation could be controlled and that phytoplankton community was structured by oyster farming. Here, ten cruises were conducted in two oyster farms (OFs) and control areas (CAs) adjacent to the Ninghai Power Plant in the upper section of Xiangshan Bay during 2009-2015. We found that thermal discharge induced severe winter algal blooms. Phytoplankton abundance and chlorophyll a (chla) were significantly lower (46.3% and 28.3%, respectively) in OF than in CA, indicating a high filtration efficiency by oysters and the associated biofouling assemblages. In addition, oyster farming significantly increased species richness (by 26.3%), Shannon-Wiener diversity (by 38.3%), and Pielou's evenness indices (by 28.8%) and reduced suspended solids (by 12.2%), total organic carbon (by 18.4%), dissolved inorganic nitrogen (by 1.5%), and phosphorus (by 3.7%). Furthermore, oyster farming considerably reduced (increased) micro-chla contribution (pheophytin/chla) by 34.8% (71.1%), suggesting a strong size-selective grazing on phytoplankton. Analysis of similarity revealed a significant difference in phytoplankton community composition between OF and CA. However, after the removal of culture rafts, all the abundance, chla, species diversity, dominant species, size structure, and community composition of phytoplankton showed no significant difference. Our study demonstrated that oyster farming effectively alleviated eutrophication and algal bloom and enhanced phytoplankton diversity, which provides guidance for aquaculture and ecological restoration in subtropical coastal eutrophic waters.

The impacts of fish cage culture and suspended oyster culture on macrobenthic communities were investigated in Xiangshan Bay, China, on a seasonal basis from January to October of 2015. Samples were collected from a fish cage farm, a suspended oyster farm, and two corresponding reference sites. Two-way ANOVA results showed that species richness, abundance, biomass, and Shannon-Wiener diversity differed significantly between the four different investigated areas, and different seasons as well. Cluster analysis showed that macrobenthic community composition in the fish and oyster culture areas significantly differed from that in the reference sites, respectively. Trophic structure of macrobenthos in the fish and oyster culture areas mostly clustered together owing to higher abundance and biomass of surface-deposit feeders and carnivores. The macrobenthic communities and feeding guilds correlated highly with environmental factors, such as bottom water chlorophyll a and nutrients, as well as sediment total organic carbon. Although integrated multi-trophic aquaculture is regarded as a suitable approach to coordinate desirable economic, social, and environmentally sustainable outcomes, coastal ecosystems may suffer from large-scale nutrient enrichment due to aquaculture and other human activities, which should not be considered in isolation.

Biogeochemical processes in the East China Sea are substantially affected by anthropogenic nutrient inputs. The dramatic decadal changes in nutrient concentrations were mainly due to the increases of DIN and DIP in the Changjiang (Yangtze) River since 1980s. As a result, phytoplankton abundance increased dramatically between 1958 and 2016 in both the Changjiang Estuary and the East China Sea. Before 1980s, chain-forming diatoms were dominant, while increasing of large-cell dinoflagellates is probably related to increasing DIN/silicate ratio. Increasing nutrient input and phytoplankton abundance greatly impact seasonal hypoxia condition in the East China Sea. Hypoxia is relatively sporadic and patchy before 2013. In 2016 and 2017, hypoxic events were more severe, occurring over larger areas with dramatically lower minimum values of dissolved oxygen. Notwithstanding, occurrences of bottom hypoxia in the East China Sea are highly dynamic and are significantly influenced by episodic events such as wind mixing.

Macrobenthos samples were collected from the Yellow and East China Seas in four seasons during 2011 to 2012. The seasonal distribution of macrobenthos and its relationship with environmental factors were analyzed. A total of 562 macrobenthic species were identified, with polychaetes and mollusks accounting for 67% of the total number of species. A similarity percentage (SIMPER) analysis showed that the dominant species were bivalve mollusks in the Yellow Sea and small-sized polychaetes in the East China Sea. A two-factor analysis of variance showed significant seasonal variations in species number, density and diversity index, and significant regional differences of biomass and density. Two-factor community similarity analysis also showed significant seasonal and regional differences in macrobenthic communities. Canonical correspondence analysis indicated that the main environmental factors affecting the macrobenthic communities were water depth, temperature, dissolved oxygen, and inorganic nitrogen. The results demonstrate significant regional differences and seasonal variations in macrobenthos in the two seas. Sediment properties and water mass characteristics are speculated to be the causes of regional differences.

Summary To examine the distribution of Trichodesmium relative to physicochemical factors during summer in the Changjiang (Yangtze River) Estuary and adjacent East China Sea shelf, three cruises were conducted separately in June 2009, July 2011, and August 2009. Trichodesmium species found were T. thiebautii , T. erythraeum , and T. hildebrandtii . The population was dominated by T. thiebautii , which accounted for >85% of the samples found. Most of them were free trichomes. Colonial forms were rarely observed (approximately 10% of our samples), occurring only in offshore waters. The depth integrated abundances of Trichodesmium were 308 × 10 3 , 1709 × 10 3 , and 3448 × 10 3 trichomes m −2 in June, July, and August, respectively. Trichodesmium was distributed abundantly in the southern or southeastern part of our study area, where nutrients were low and light penetration, temperature, and salinity were high, which were influenced by the Taiwan Warm Current (TWC) and Kuroshio. Trichodesmium was found in low abundance in inshore, eutrophic, low-salinity waters, which were mainly controlled by the Changjiang Diluted Water (CDW) and coastal current. These results suggest that spatiotemporal changes in the summer Trichodesmium distribution correlate highly with the variations in physicochemical properties that are primarily controlled by the TWC, Kuroshio, and CDW. The summer N 2 fixation rate of Trichodesmium was estimated at 12.3 μmol N m −2 d −1 in our study area, contributing >50% of biological N 2 fixation.

To evaluate the effects of different anthropogenic activities on zooplankton and the pelagic ecosystem, we conducted seasonal cruises in 2010 to assess spatial heterogeneity among the mesozooplankton communities of Xiangshan Bay, a subtropical semi-enclosed bay in China. The evaluation included five different areas: a kelp farm, an oyster farm, a fish farm, the thermal discharge area of a power plant, and an artificial reef, and we aimed to identify whether anthropogenic activities dominated spatial variation in the mesozooplankton communities. The results demonstrated clear spatial heterogeneity among the mesozooplankton communities of the studied areas, dominantly driven by natural hydrographic properties, except in the area near the thermal discharge outlet of the power station. In the outlet area, thermal shock caused by the discharge influenced the mesozooplankton community by decreasing abundance and biomass throughout the four seasons, even causing a shift in the dominant species near the outlet during summer fromAcartia pacificato eurythermal and warm water taxa. Unique features of the mesozooplankton community in the oyster farm may be due to the combined effects of oyster culture and the natural environment in the branch harbour. However, kelp and fish culture, and the construction of an artificial reef did not exert any obvious influence on the mesozooplankton communities up to 2010, probably because of the small scale of the aquaculture and a time lag in the rehabilitation effects of the artificial reef. Thus, our results suggested that the dominant factors influencing spatial variations of mesozooplankton communities in Xiangshan Bay were still the natural hydrographic properties, but the thermal discharge was an anthropogenic activity that changed the pelagic ecosystem, and should be supervised.

Coastal eutrophication and its associated harmful algal blooms have emerged as one of the most severe environmental problems worldwide. Seaweed cultivation has been widely encouraged to control eutrophication and algal blooms. Among them, cultivated kelp (Saccharina japonica) dominates primarily by production and area. However, the responses of water quality and phytoplankton community to kelp farming remain unclear. Here, thirteen cruises were conducted in the kelp farms and control areas in the turbid, highly eutrophic Xiangshan Bay of the East China Sea from 2008 to 2015. Results indicated that kelp cultivation slightly increased dissolved oxygen and pH, but reduced dissolved inorganic nitrogen and phosphorus. We estimated that kelp harvesting would remove 297 t of nitrogen and 42 t of phosphorus from this bay annually. Because of decreased flow velocity, turbulence, and sediment resuspension, kelp farming greatly reduced suspended solids and increased transparency, resulting in increases in phytoplankton chlorophyll a and abundance. Additionally, kelp farming appreciably increased phytoplankton species number, Marglef richness, and Shannon–Wiener diversity indices by 51.6%, 40.1%, and 13.1%, respectively. Analysis of similarity and similarity percentages demonstrated that phytoplankton community composition differed significantly between the farm and control area, which was mostly attributed to long-chained diatoms and single-celled dinoflagellates. However, after the kelp harvesting, all measurements of water quality and phytoplankton biomass, diversity, and community composition exhibited no significant difference. Our study highlights that kelp cultivation alleviates eutrophication and acidification and enhances phytoplankton diversity, thus providing guidance for macroalgal aquaculture and remediation in eutrophic waters.

To explore the effects of coastal eutrophication and warming on phytoplankton biomass and cell size, we analyzed current and historical data for size-fractionated chlorophyll a (chla) in Xiangshan Bay, China. Results showed that micro- and nanophytoplankton overwhelmingly dominated (>84%) in all seasons. The contribution of micro-chla was significantly lower in warm than in cold seasons, whereas contribution of pico-chla showed the opposite result. Overall, the micro-chla contribution increased with decreasing pico-chla contribution from the stable, clear, eutrophic upper bay to the turbulent, turbid lower bay, indicating that phytoplankton size structure on a spatial scale was largely shaped by water column stability and light rather than by nutrients. Since the 1980s, phytoplankton biomass, primary productivity, and micro-chla contribution in Xiangshan Bay have increased sharply with increasing nutrient amounts and temperature. Additionally, it seems that algal bloom seasonality has shifted forward from spring to winter since the power plant operations in 2006.

Benthic macrofaunal communities along an estuarine gradient of a macrotidal estuary of the Jiaojiang River, China were analyzed to reveal spatial variations in those communities and their relationships with environmental variables. Benthic macrofauna were collected from 15 sampling stations in May and October 2010. Polychaetes and Mollusks are the dominant taxonomic groups. The species richness, density and Shannon diversity index generally tended to increase as one moved away from shore toward the outer stations of the estuary. Cluster analysis and multidimensional scaling ordinations indicated the presence of three distinct zones of macrofaunal communities in the estuary. Multivariate analyses showed a strong relationship exists between the macrofaunal communities and the environmental parameters in the Jiaojiang River estuary. Aside from salinity, dissolved inorganic nitrogen and soluble reactive phosphate contents were also the major factor affecting the structure of the macrofaunal communities. The results demonstrated the important effects of nutrient-load on the distributional patterns of benthic macrofaunal communities in nutrient-rich estuaries.

To explore the distribution and composition of phytoplankton community and their responses to environmental changes, summer net-collected phytoplankton and physicochemical parameters in the Hangzhou Bay during 2004–2010 were investigated. A total of four phyla and 84 species were identified, including 67 diatom and 12 dinoflagellate species. The dominant species constantly consisted of the diatoms, although the dominance of dinoflagellate and cyanobacteria increased recently. Due to great spatio-temporal variations in environmental factors (salinity, suspended solids, and nutrient concentration), significant heterogeneities in community compositions among different years and subregions (inner and middle sections, and bay mouth) were found based on the analyses of multidimensional scaling and similarity. Canonical correspondence analysis showed that salinity and Si/N were the main variables associated with algal assemblage. Compared with the historical data since the 1980s, eutrophication (N, P, and N/P increased with decreasing Si/N) was exacerbated drastically. Moreover, climatic forcing and human activities resulted in a series of physical alterations, including sediment retention, temperature increase, and salinity decrease as well as reduction in water exchanges. All these changes induced obvious increases in cell density and Chl-a while decreases in species diversity and diatom-dinoflagellate ratio as well as the shifting of dominant species. Therefore, the long-term phytoplankton variations were closely related to anthropogenic and climatic perturbations in the Hangzhou Bay.

Quantitative and precise observation of the phytoplankton is required for biogeochemical and environmental monitoring in coastal seas. However, phytoplankton data may vary greatly depending on the sampling methods. Net and water collection methods are widely adopted for obtaining marine phytoplankton in China; yet, exact differences and relationships between water- and net-collected phytoplankton communities remain poorly understood. Here, annual phytoplankton surveys in the southern Yellow Sea were conducted using these two sampling methods. Compared with water collection, net collection resulted in the loss of numerous noncolonial small-celled species, particularly cryptophytes and coccolithophores, and reduced phytoplankton depth-averaged abundance (DAA) by 1–2 orders of magnitude. However, through net collection, some scarce, large, or colonial species with low abundances were obtained, and the net-collected DAA distribution was highly consistent with the water-collected DAA distribution in each season. Moreover, DAAs of planktonic large-celled or long-chained dominant species, belonging to diatom, dinoflagellate, and cyanobacteria, were comparable in net- and water-collected samples. Regression analysis showed highly significant regression coefficients between net- and water-collected DAAs of phytoplankton, diatom, and diatom plus dinoflagellate, suggesting that net-collected data could be roughly derived from water-collected results. Non-metric multidimensional scaling and analysis of similarity indicated that phytoplankton community composition significantly differed between net- and water-collected samples. Similarity percentages confirmed that this difference was largely attributed to small-sized benthic and meroplanktonic diatoms, unidentified cryptophytes and coccolithophores, and single-celled dinoflagellates. Our results suggest that the sampling method profoundly influences phytoplankton analysis in terms of species information, abundance, dominant species, and community composition.

We analyzed the composition and distribution of phytoplankton in relation to physicochemical factors in the Changjiang (Yangtze River) Estuary and adjacent East China Sea shelf in June and August 2009. Diatoms and dinoflagellates dominated the community, particularly in eutrophic inshore waters controlled by the Changjiang Diluted Water (CDW), coastal current, and upwelling. However, high densities of cyanobacteria and cryptophytes were observed in the oligotrophic offshore waters influenced by the Taiwan Warm Current (TWC) and Kuroshio. In June, the northeastward CDW plume combined with the Yellow Sea Coastal Current induced algal bloom in the northern part of the CE. In August, the enhanced CDW formed two narrow, low-salinity tongues that extended eastward and southward (associated with the upwelling and coastal current), resulting in phytoplankton blooms off the CE and in the Zhejiang coastal waters, respectively. Phytoplankton abundance in August was considerably higher than in June, with increased solar radiation, CDW, and upwelling. The maximum abundance occurred on the surface in inshore turbid waters and on the subsurface (5–30 m) in offshore clear waters with increased stratification. Based on multidimensional scaling and cluster analysis, we found appreciable spatio-temporal variations in algal community composition. Different ecological groups corresponded with hydrographic distributions. Canonical correspondence analysis showed that nutrients, salinity, temperature, and suspended particulate matter were the main variables associated with community distribution. We suggest that the variations in summer phytoplankton community are highly correlated with the significant monthly and spatial variability in physicochemical properties, which are primarily controlled by the CDW and TWC.

The distributions of N utilizing bacteria (denitrifying bacteria and ammonifying bacteria), P utilizing bacteria (organic phosphobacteria and inorganic phosphobacteria) and heterotrophic bacteria in the Changjiang Estuary, and the roles of main environmental factors in distributing bacteria, are explored with observations from two cruises in June and August 2006. Comparisons between the two important periods of initial hypoxia phase (June) and developed hypoxia phase (August) show differences in both bacterial distributions and the associated main environmental factors. First, the primary group of ammonifying bacteria has larger magnitude with spatial maximum value in the hypoxic stations related to sediment in August. The phosphobacterial abundance and detection rates in August are much lower than those in June, but the denitrifying bacterial abundance becomes greater in August. However, the difference of heterotrophic bacterial abundance between June and August is not obvious. Second, main environmental factors influencing bacteria vary from initial hypoxia phase to developed hypoxia phase. Two parameters (salinity and NO3−) in surface water and five environmental parameters (pH, salinity, PO43−, NO3− and temperature) in bottom water and surface sediment play major roles in the bacterial abundance in June, while different parameter combinations (salinity and PO4−) in surface water and different parameter combinations (DO, DOC, NO3−, PO43− and pH) in bottom water and surface sediment play major roles in August. Moreover, the bottom bacteria distributions in area south of 31°N are related to the position of the Taiwan Warm Current in June. The bacterial abundance and distribution may respond to the environmental change in the hypoxia processes of initial phase and developed phase. During the hypoxia processes, the whole structure of bacterial functional groups probably turns to different states, causing the recycling of nutrient regeneration and aggravating hypoxia regionally.

There is concern about the impact of expanding coastal aquaculture activity, particu- larly in China, with the biggest aquaculture industry in the world. We conducted 3 cruises to the oyster, kelp and fish farms of the Xiangshan Bay during the winterspring transition (02/2009), winter (01/2010) and spring (04/2010). Our aim was to explore the spatial heterogeneity of the phytoplankton community in different culture habitats and evaluate the effects of mariculture on phytoplankton. Different culture types formed unique small-scale habitats that exhibited various en vironmental gradients and forms of phytoplankton community patchiness. Both environmental (i.e. temperature, salinity, transparency, suspended solids, dissolved oxygen, pH, dissolved inor- ganic nitrogen, PO4-P, SiO3-Si, N/P, and total organic carbon) and phytoplankton (i.e. cell density, chlorophyll a, species richness, main dominant species, and community structure) parameters were significantly different among the 3 culture habitats. Canonical correspondence analysis showed that phytoplankton distribution was mainly influenced by temperature, nutrition, salinity and suspended solids. Oyster and kelp farming effectively alleviated coastal eutrophication and also increased species richness and diversity; fish farming did not. Temperature elevation caused by the thermal discharge from a power plant induced phytoplankton blooms during the winter and winterspring transition. Nevertheless, the filtering effect of suspended oysters dramatically reduced microalgal biomass. The kelp and oyster farms were much better than the fish farm in terms of water quality, phytoplankton composition, and habitat restoration function. These results provide guidelines for future policy-making in aquaculture as well as eutrophication control and management.

The phytoplankton community in the Changjiang Estuary is subject to intensive physical and chemical stresses because of human- and climate-driven changes. We obtained historical data on summer phytoplankton communities from 1959 to 2009 to explore responses to long-term environmental changes. The nitrogen (N) and phosphorus (P) concentrations and ratios (N/P and N/Si) increased because of persistent riverine loading, but silicon (Si) levels remained constant. Climatic changes and extensive water diversions and withdrawals (sediment retention) resulted in a series of physical alterations, including increased temperature, turbidity reduction, and river plume shrinkage. These changes induced a dramatic increase in microalgal biomass (cell density and chlorophyll a) with a decreasing diatom-dinoflagellate ratio, and exacerbated harmful algal blooms. In the past dominant net-collected species were usually chain-forming diatoms; however, more recently, large dinoflagellates and filamentous cyanobacteria dominate. This was not consistent with information from water samples (co-dominated by small dinoflagellates), because of the loss of solitary species collected using a 76-μm net. The dominant species shifted from temperate-subtropical/eurythermal to subtropical-tropical/eurythermal taxa in the warmer water caused by global warming and hydrographic changes. There was also an increased dominance of euryhaline/high-salinity species due to increase in Kuroshio transport and the northward Taiwan Warm Current and reduction in Changjiang Diluted Water. All these changes in phytoplankton communities appear to be closely related to an increase in anthropogenic activities and climatic changes.

Six cruises were conducted in a fish farm adjacent to the Ninghai Power Plant in Xiangshan Bay, East China Sea. Fish farming significantly increased NH 4 + , DIP, and TOC concentrations, while it significantly decreased the DO level. These increase/decrease trends were more pronounced in warmer seasons. Although culture practices did not significantly increase phytoplankton density, it drastically enhanced dinoflagellate abundance and domination. Significant differences in species diversity and community composition between the cages and the control area were also observed. Temperature elevation caused by thermal discharge associated with eutrophication resulted in a dominant species shift from diatoms alone to dinoflagellates and diatoms. This is the first report of stress-induced toxic dinoflagellate (Prorocentrum minimum) blooms in winter and the winter–spring transition in this bay. Therefore, the effects of aquaculture activity and power plant construction in such a eutrophic, semi-enclosed bay require further attention.