黄、东海浮游植物功能群研究
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摘要
浮游植物是海洋中最主要的初级生产者,启动了海洋生态系统的能量流和物质流,开启了海洋食物链。973项目“我国近海生态系统食物产出的关键过程及其可持续机理”提出开展从食物网的原核生物-浮游植物到顶级生物从个体到粒径谱、功能群和营养层的全程(from end to end)食物网研究,并将生物功能群的食物网营养动力学作为拟解决的关键科学问题,因此需要对浮游植物功能群的组成和生态特征进行研究。
本论文基于2006年4月至2008年6月在黄、东海进行的9次多学科综合外业调查,利用浮游植物群落资料,研究了该海域浮游植物功能群的组成及生物量的时空分布特征,分析了浮游植物功能群与环境因子的关系、以及关键物理过程对浮游植物功能群的影响。
根据营养功能将浮游植物划分为12个功能群:微微型浮游植物(PicoPl)、微型硅藻(NanDiat)、微型自养鞭毛藻(NanAutFl)、小型自养甲藻(MAutFl)、小型混合营养甲藻(MMixFl)、小型异养甲藻(MHetFl)、小型单体硅藻(MUniDiat)、小型无刺群体硅藻(MHairDiat)、小型有刺群体硅藻(MBranDiat)、产毒浮游植物(ToxPl)、不可食浮游植物(InedPl)、其它浮游植物(Others)。
浮游植物水柱生物量在南黄海春季高于秋季;春季高值区位于山东半岛近岸水域,秋季高值区位于济州岛附近水域。在长江口和东海水域夏季最高,春季次之,秋季较低,冬季最低;高值区多分布在长江口-浙江近岸水域,外海高盐区生物量较低。在垂直方向上,浮游植物生物量主要分布在浅层水体。
黄、东海浮游植物功能群组成时空变化较大:南黄海春、秋季MHairDiat都是最优势的功能群;长江口邻近黄东海,春末以MHetFl为主的甲藻功能群在群落占优,ToxPl有较高的比例;夏中,硅藻取代甲藻,MHairDiat成为最优势功能群,MBranDiat也有较高的比例;夏末,硅藻功能群占群落比例超过95%;东海陆架区秋冬季均以硅藻功能群为主,MHairDiat > MUniDiat > MBranDiat;长江口及东海近岸区甲藻功能群占优,5月份MMixFl > MAutFl > MHetFl,6月份MHetFl取代MMixFl成为最优势功能群。
MHairDiat春季在山东半岛东端的海槽区出现全年最高值;春末到夏末,长江口及东海近岸区水柱生物量逐渐升高;生物量从秋季到冬季持续下降,秋季黄海冷水团水域生物量极低。MUniDiat在春季的分布与MHairDiat类似;从夏中至夏末,长江口及东海近岸区生物量持续升高;秋季,生物量在东海陆架区高于南黄海;冬季,东海陆架区生物量略有下降,在长江口附近形成高值区。MBranDiat春季在长江口邻近黄海生物量较高,在东海少有分布,春末开始东海生物量升高,高值区分布在杭州湾外;秋季生物量下降,在黄海分布在近岸,东海分布在外海;冬季生物量低、分布范围小。MHetFl春季在山东半岛近岸、长江口-杭州湾外有很高的生物量;夏季开始,生物量显著降低,冬季仅在东海陆架外缘有分布。MMixFl春季在东海高于南黄海,春末生物量在长江口及东海近岸区上升,夏季开始降低;秋季在南黄海的分布与春季类似,高于东海;冬季,东海陆架区生物量很低。MAutFl春末在长江口邻近黄海出现全年最高值,高生物量从夏季开始衰退,秋、冬季生物量均很低。ToxPl在春夏季生物量较高,春季主要分布在南黄海中部、长江口和东海近岸区;夏中长江口邻近黄海出现全年最大值,到夏末消失。InedPl夏末在长江口邻近黄海出现全年最高值,其生物量在外海高盐水域较高。
浮游植物关键种的演替明显:南黄海关键种主要为硅藻,春季为太平洋海链藻(Thalassiosira pacifica)、豪猪棘冠藻(Corethrom hystrix)等,秋季为斯氏几内亚藻(Guinardia striata)、圆筛藻(Coscinodiscus sp.)。长江口及东海水域,5、6月份关键种以甲藻为主,夜光藻(Noctiluca scientillans)、具齿原甲藻(Prorocentrum dentatum)、米氏凯伦藻(Karenia mikimotoi)等水华物种、柔弱伪菱形藻(Pseudo-nitzschia delicatissima)等产毒物种都有较高的生物量;夏季,甲藻式微,脆指管藻(Dactyliosolen fragilissimus)、旋链角毛藻(Chaetoceros curvisetus)、厚刺根管藻(Rhizosolenia crassipina)和环纹劳德藻(Lauderia annulata)等硅藻成为关键种;秋季,关键种为圆筛藻、圆海链藻(Thalassiosira rotula)、斯氏几内亚藻(Guinardia striata)等硅藻;冬季,关键种主要是圆筛藻等单体硅藻。夜光藻、具齿原甲藻、米氏凯伦藻春季在长江口附近水域易形成水华(赤潮),而太平洋海链藻等硅藻春季在南黄海也有爆发水华的可能。
浮游植物功能群与环境因子的关系存在时空差异:MHairDiat在黄东海的分布主要受温度和盐度的影响,但春季在黄海、夏季在长江口及邻近黄东海可能受到营养盐的限制;MUniDiat的分布主要受营养盐影响,生物量在高营养盐水体较高,春季在南黄海主要受盐度影响;MBranDiat在春、夏季可能受营养盐限制,在夏末、秋季和冬季的分布主要受温度和盐度影响;MAutFl的分布春季在南黄海受磷酸盐影响,在长江口受铵盐影响,秋季在东海陆架区与营养盐密切相关,其它航次主要受温度和盐度影响;MMixFl的分布主要受温度或/和盐度的影响,春季在南黄海、秋季在长江口及东海陆架区的分布主要受氮盐的影响;MHetFl主要分布在近岸的表层水体,在外海高盐高温水体也能形成较高的生物量;ToxPl的分布与营养盐的关系在秋季不明显,春季在南黄海、夏季在长江口、冬季在东海陆架区主要受氮盐的影响;InedPl的分布主要受温度和盐度的影响,但秋、冬季在东海陆架区在高营养盐水体分布较多。
黄海冷水团水域浮游植物生物量极低,温跃层的存在阻碍了营养盐向上输送,表层受营养盐特别是磷酸盐限制,仅适宜异养甲藻;底层受光限制,仅适宜底栖硅藻。长江冲淡水携带的大量营养盐支持了长江口水域的高生物量,生物量锋面出现在光和营养盐取得最佳权衡的水域。高盐、低营养盐的黑潮和台湾暖流控制区域,浮游植物难以形成高生物量。
综合基于浮游植物功能群组成的聚类分析结果和群落生物量对黄、东海进行了生态区的划分:南黄海春季划分为4个生态区、秋季划分为3个生态区,长江口及邻近黄东海水域春末划分为2个生态区、夏季划分为4个生态区、夏末划分为3个生态区,东海陆架区秋、冬季均划分为2个生态区,长江口及东海近岸区春季划分为2个生态区、春末划分为3个生态区。
春、秋季,基于浮游植物功能群对南黄海生态区的划分与基于浮游动物功能群的结果类似,但生物量分布模式不同;浮游植物功能群与中华哲水蚤的摄食、代谢、生殖行为有密切的关系。夏季,长江口外高生物量的MHairDiat和MBranDiat与底层的氧亏损有关。
本文从生物量出发,报道了黄、东海浮游植物功能群、关键种的组成及时空分布,研究了各功能群与环境因子的关系,并基于浮游植物功能群对黄、东海进行了生态区的划分,这些结果为黄、东海食物产出关键过程的预测模型提供了基础数据。今后的研究需要综合多种分析方法,获取全粒径浮游植物资料,以全面分析浮游植物功能群结构;同时与高营养层次生物功能群进行整合研究,以更好的理解中国近海食物网的营养结构。
Phytoplankton is the main component of primary producers in the ocean, which triggers the energy flow and material flow of marine ecosystem, and initiates the pelagic food web. Food web from end to end is the core target of 973 key project―Key processes and sustainable mechanism of food output in Chinese coastal pelagic ecosystem‖. Trophic dynamics of plankton functional groups is one of the key scientific issues. Therefore, it is critical to study the composition and ecological characteristics of phytoplankton functional groups (PFGs) in the Yellow Sea (YS) and the East China Sea (ECS).
From April 2006 to June 2008, nine cruises were conducted in the YS and the ECS. Based on phytoplankton data by Unterm?hl method, composition and ecological characteristics of PFGs were investigated. Relationships between PFGs and environmental factors were analyzed by CCA, and influences of hydrodynamic processes on PFGs were taken in account as well.
According to trophic function, phytoplankton is divided into 12 functional groups, i.e. pico-phytoplankton (PicoPl), nano-diatom (NanDiat), autotrophic nano-flagellate (NanAutFl), autotrophic micro-dinoflagellate (MAutFl), mixotrophic micro-dinoflagellate (MMixFl), heterotrophic micro-dinoflagellate (MHetFl), unicellular micro-diatom (MUniDiat), hair-shaped micro-diatom (MHairDiat), branch-shaped micro-diatom (MBranDiat), toxic phytoplankton (ToxPl), inedible phytoplankton (InedPl), and other phytoplankton (Others).
Phytoplankton integrated biomass in spring was much higher than that in autumn in the southern YS. High biomass distributed in the coastal water of Shandong Peninsula in spring, whereas, in the coastal water of Cheju Island in autumn. Phytoplankton integrated biomass showed evident seasonal variation in the Yangtze River Estuary (YRE) and its adjacent waters, i.e., in summer > in spring > in autumn > in winter. Biomass in the coastal waters of YRE and Zhejiang Province was much higher than that in off-shore sites with salinity > 31. Vertically, phytoplankton biomass decreased with depth from surface to bottom.
Composition of PFGs varied spatially and temporally in the YS and the ECS. MHairDiat was the most dominant group in the southern YS in spring and autumn. MHetFl-dominant dinoflagellate groups were most important, in addition, ToxPl was also important in the YRE and its adjacent waters in late spring. Diatom groups replaced dinoflagellate, and MHairDiat and MBranDiat were dominant in the YRE and its adjacent waters in the middle of summer. Diatom groups accounted for 95% of phytoplankton biomass in the YRE and its adjacent waters in late summer. Diatom groups, MHairDiat > MUniDiat > MBranDiat, were dominant in the continental shelf water of ECS in autumn and winter. Dinoflagellate groups, MMixFl > MAutFl > MHetFl, were dominant in the YRE and the coastal ECS in May of 2007, while MHetFl > MMixFl > MAutFl in June of 2008.
MHairDiat was most abundant in the YS Trough water near Shandong Peninsula in spring. Its biomass in the YRE and the coastal ECS increased from late spring to late summer, and decreased from autumn to winter. MHairDiat showed extremely low biomass in the Yellow Sea Cold Water Mass (YSCWM) area in autumn. Distribution of MUniDiat was similar to that of MHairDiat in spring. Its biomass in the YRE and the coastal ECS increased from August to October. MUniDiat was more abundant in the ECS than in the southern YS in autumn. MUniDiat biomass in winter was lower than that in autumn in the ECS, and was densely distributed near the YRE. In spring, MBranDiat was abundant in the YS water near the YRE, while scarce in the ECS. MBranDiat biomass increased from June to October, and decreased from autumn to winter in the ECS. Late spring peak value appeared in coastal water of Hangzhou Bay, while autumn peak value in the open sea. In autumn MBranDiat was densely distributed in the coastal water of YS. High density of MHetFl located in coastal water of Shandong Peninsula and the area near the YRE and Hangzhou Bay in spring. MHetFl biomass was much low in the YS and the ECS from summer to winter. MMixFl biomass was highest in late spring among four seasons, and its peak value occurred in the YRE and the coastal ECS. MMixFl biomass was higher in the ECS than in the YS in spring, while lower in the ECS than in the YS in autumn. MMixFl biomass was extremely low in the continental shelf water of ECS in winter. Maximum MAutFl biomass occurred in the YRE adjacent YS in late spring, and high biomass decreased in summer. MAutFl was in low abundance in the continental shelf water of ECS in autumn and winter. ToxPl was abundant in the middle of southern YS, in the YRE and the coastal ECS in spring, and showed peak biomass, which disappeared in late summer, in the YRE adjacent YS in the middle of summer. InedPl was most abundant in the YRE adjacent YS in late summer. Generally, its biomass was high in open sea.
Succession of keystone species was evident in survey area. Keystone species were mainly diatoms in the southern YS. Keystone species were Thalassiosira pacifica and Corethrom hystrix in spring, and Guinardia striata and Coscinodiscus sp. in autumn. In the YRE and the ECS, dinoflagellates, such as Noctiluca scientillans, Prorocentrum dentatum, Karenia mikimotoi, became keystone species in May and June, besides toxic phytoplankton Pseudo-nitzschia delicatissima showed high biomass as well. Diatom, i.e. Dactyliosolen fragilissimus, Chaetoceros curvisetus, Rhizosolenia crassipina, Lauderia annulata, replaced dinoflagellate and became keystone species in summer. Diatoms, such as Coscinodiscus sp., Thalassiosira rotula, Guinardia striata, became keystone species again in autumn. Keystone species in winter was mainly unicellular diatom, such as Coscinodiscus sp.. Noctiluca scientillans, Prorocentrum dentatum, and Karenia mikimotoi could bloom in the YRE and its adjacent water in spring, so could Thalassiosira pacifica in the southern YS in spring.
Correlations between PFGs and environmental factors were highly variable in spatial and temporal scale. MHairDiat was well correlated with salinity and temperature in survey area, however, it could be limited by nutrients during cruise LST-I and cruise QYQ-II. Distributions of MUniDiat were mainly influenced by nutrients, but also correlated with salinity during cruise LST-I. Distributions of MBranDiat could be influenced by nutrients in spring and summer, while by temperature and salinity in late summer, autumn and winter. Phosphate was the key factor that influenced the distributions of MAutFl during cruise LST-I, while ammonia in the Yangtze River Estuary, and nutrients during cruise ZLJ-I, and temperature and salinity during other cruises. MMixFl well correlated with temperature and salinity, whereas correlated with nitrogen during cruise LST-I and cruise ZLJ-I. MHetFl mainly distributed in upper layers of coastal waters, however, it could show high biomass in open sea as well. ToxPl was mainly influenced by nitrogen during cruise LST-I, cruise QYQ-II, and cruise ZLJ-II. Distributions of InedPl were mostly controlled by temperature and salinity, in addition, InedPl positively correlated with nutrient during cruise ZLJ-I and cruise ZLJ-II.
Phytoplankton biomass was extremely low in the YSCWM in autumn, where the strong thermocline stopped the upward transport of nutrients from bottom water. There was mainly phosphate limitation in surface water, where MMixFl and MHetFl were dominant, and illumination limitation below surface, where MUniDiat was dominant. High phytoplankton biomass in the YRE was supported by rich nutrients transported by Yangtze River Dilute Water. Phytoplankton biomass front appeared in the waters where nutrients and illumination stroke a balance. Phytoplankton biomass was generally low in Kuroshio and Taiwan Warm Current, characteristic of high salinity and low nutrients.
Cluster analysis of survey sites of every cruise was conducted based on PFGs composition. According to cluster analysis and phytoplankton biomass, the YS and the ECS were divieded into several provinces. The southern YS were divided into 4 provinces in spring, while 3 provinces in autumn. The YRE and its adjacent YS and ECS was divided into 2 provinces in late spring, and 4 provinces in the middle of summer, and 3 provinces in late summer. The continental shelf water of ECS was divided into 2 provinces in both autumn and winter. The YRE and the coastal ECS were divided into 2 provinces in spring and 3 provinces in late spring.
PFGs provinces were similar to zooplankton functional groups provinces in the southern YS in spring and autumn, however, they were quiet different in distribution of biomass. Biomass and composition of PFGs was closely correlated with grazing, metabolism, and reproduction of Calanus sinicus. High biomass of MHairDiat and MBranDiat could be a key factor in formation of hypoxia in the YRE in summer.
Based on carbon biomass, the composition and distributions of PFGs were reported in the YS and the ECS, so was the distribution of keystone species. Correlations between PFGs and environmental factors were analyzed, and PFGs provinces were divided as well. This dissertation provides for the food output model in the YS and the ECS. To understand the PFGs comprehensively, multi-method should be employed in the analysis of phytoplankton samples. Furthermore, interdisciplinarity study should be conducted to get insights into trophic structure of pelagic food web in the YS and the ECS.
本论文基于2006年4月至2008年6月在黄、东海进行的9次多学科综合外业调查,利用浮游植物群落资料,研究了该海域浮游植物功能群的组成及生物量的时空分布特征,分析了浮游植物功能群与环境因子的关系、以及关键物理过程对浮游植物功能群的影响。
根据营养功能将浮游植物划分为12个功能群:微微型浮游植物(PicoPl)、微型硅藻(NanDiat)、微型自养鞭毛藻(NanAutFl)、小型自养甲藻(MAutFl)、小型混合营养甲藻(MMixFl)、小型异养甲藻(MHetFl)、小型单体硅藻(MUniDiat)、小型无刺群体硅藻(MHairDiat)、小型有刺群体硅藻(MBranDiat)、产毒浮游植物(ToxPl)、不可食浮游植物(InedPl)、其它浮游植物(Others)。
浮游植物水柱生物量在南黄海春季高于秋季;春季高值区位于山东半岛近岸水域,秋季高值区位于济州岛附近水域。在长江口和东海水域夏季最高,春季次之,秋季较低,冬季最低;高值区多分布在长江口-浙江近岸水域,外海高盐区生物量较低。在垂直方向上,浮游植物生物量主要分布在浅层水体。
黄、东海浮游植物功能群组成时空变化较大:南黄海春、秋季MHairDiat都是最优势的功能群;长江口邻近黄东海,春末以MHetFl为主的甲藻功能群在群落占优,ToxPl有较高的比例;夏中,硅藻取代甲藻,MHairDiat成为最优势功能群,MBranDiat也有较高的比例;夏末,硅藻功能群占群落比例超过95%;东海陆架区秋冬季均以硅藻功能群为主,MHairDiat > MUniDiat > MBranDiat;长江口及东海近岸区甲藻功能群占优,5月份MMixFl > MAutFl > MHetFl,6月份MHetFl取代MMixFl成为最优势功能群。
MHairDiat春季在山东半岛东端的海槽区出现全年最高值;春末到夏末,长江口及东海近岸区水柱生物量逐渐升高;生物量从秋季到冬季持续下降,秋季黄海冷水团水域生物量极低。MUniDiat在春季的分布与MHairDiat类似;从夏中至夏末,长江口及东海近岸区生物量持续升高;秋季,生物量在东海陆架区高于南黄海;冬季,东海陆架区生物量略有下降,在长江口附近形成高值区。MBranDiat春季在长江口邻近黄海生物量较高,在东海少有分布,春末开始东海生物量升高,高值区分布在杭州湾外;秋季生物量下降,在黄海分布在近岸,东海分布在外海;冬季生物量低、分布范围小。MHetFl春季在山东半岛近岸、长江口-杭州湾外有很高的生物量;夏季开始,生物量显著降低,冬季仅在东海陆架外缘有分布。MMixFl春季在东海高于南黄海,春末生物量在长江口及东海近岸区上升,夏季开始降低;秋季在南黄海的分布与春季类似,高于东海;冬季,东海陆架区生物量很低。MAutFl春末在长江口邻近黄海出现全年最高值,高生物量从夏季开始衰退,秋、冬季生物量均很低。ToxPl在春夏季生物量较高,春季主要分布在南黄海中部、长江口和东海近岸区;夏中长江口邻近黄海出现全年最大值,到夏末消失。InedPl夏末在长江口邻近黄海出现全年最高值,其生物量在外海高盐水域较高。
浮游植物关键种的演替明显:南黄海关键种主要为硅藻,春季为太平洋海链藻(Thalassiosira pacifica)、豪猪棘冠藻(Corethrom hystrix)等,秋季为斯氏几内亚藻(Guinardia striata)、圆筛藻(Coscinodiscus sp.)。长江口及东海水域,5、6月份关键种以甲藻为主,夜光藻(Noctiluca scientillans)、具齿原甲藻(Prorocentrum dentatum)、米氏凯伦藻(Karenia mikimotoi)等水华物种、柔弱伪菱形藻(Pseudo-nitzschia delicatissima)等产毒物种都有较高的生物量;夏季,甲藻式微,脆指管藻(Dactyliosolen fragilissimus)、旋链角毛藻(Chaetoceros curvisetus)、厚刺根管藻(Rhizosolenia crassipina)和环纹劳德藻(Lauderia annulata)等硅藻成为关键种;秋季,关键种为圆筛藻、圆海链藻(Thalassiosira rotula)、斯氏几内亚藻(Guinardia striata)等硅藻;冬季,关键种主要是圆筛藻等单体硅藻。夜光藻、具齿原甲藻、米氏凯伦藻春季在长江口附近水域易形成水华(赤潮),而太平洋海链藻等硅藻春季在南黄海也有爆发水华的可能。
浮游植物功能群与环境因子的关系存在时空差异:MHairDiat在黄东海的分布主要受温度和盐度的影响,但春季在黄海、夏季在长江口及邻近黄东海可能受到营养盐的限制;MUniDiat的分布主要受营养盐影响,生物量在高营养盐水体较高,春季在南黄海主要受盐度影响;MBranDiat在春、夏季可能受营养盐限制,在夏末、秋季和冬季的分布主要受温度和盐度影响;MAutFl的分布春季在南黄海受磷酸盐影响,在长江口受铵盐影响,秋季在东海陆架区与营养盐密切相关,其它航次主要受温度和盐度影响;MMixFl的分布主要受温度或/和盐度的影响,春季在南黄海、秋季在长江口及东海陆架区的分布主要受氮盐的影响;MHetFl主要分布在近岸的表层水体,在外海高盐高温水体也能形成较高的生物量;ToxPl的分布与营养盐的关系在秋季不明显,春季在南黄海、夏季在长江口、冬季在东海陆架区主要受氮盐的影响;InedPl的分布主要受温度和盐度的影响,但秋、冬季在东海陆架区在高营养盐水体分布较多。
黄海冷水团水域浮游植物生物量极低,温跃层的存在阻碍了营养盐向上输送,表层受营养盐特别是磷酸盐限制,仅适宜异养甲藻;底层受光限制,仅适宜底栖硅藻。长江冲淡水携带的大量营养盐支持了长江口水域的高生物量,生物量锋面出现在光和营养盐取得最佳权衡的水域。高盐、低营养盐的黑潮和台湾暖流控制区域,浮游植物难以形成高生物量。
综合基于浮游植物功能群组成的聚类分析结果和群落生物量对黄、东海进行了生态区的划分:南黄海春季划分为4个生态区、秋季划分为3个生态区,长江口及邻近黄东海水域春末划分为2个生态区、夏季划分为4个生态区、夏末划分为3个生态区,东海陆架区秋、冬季均划分为2个生态区,长江口及东海近岸区春季划分为2个生态区、春末划分为3个生态区。
春、秋季,基于浮游植物功能群对南黄海生态区的划分与基于浮游动物功能群的结果类似,但生物量分布模式不同;浮游植物功能群与中华哲水蚤的摄食、代谢、生殖行为有密切的关系。夏季,长江口外高生物量的MHairDiat和MBranDiat与底层的氧亏损有关。
本文从生物量出发,报道了黄、东海浮游植物功能群、关键种的组成及时空分布,研究了各功能群与环境因子的关系,并基于浮游植物功能群对黄、东海进行了生态区的划分,这些结果为黄、东海食物产出关键过程的预测模型提供了基础数据。今后的研究需要综合多种分析方法,获取全粒径浮游植物资料,以全面分析浮游植物功能群结构;同时与高营养层次生物功能群进行整合研究,以更好的理解中国近海食物网的营养结构。
Phytoplankton is the main component of primary producers in the ocean, which triggers the energy flow and material flow of marine ecosystem, and initiates the pelagic food web. Food web from end to end is the core target of 973 key project―Key processes and sustainable mechanism of food output in Chinese coastal pelagic ecosystem‖. Trophic dynamics of plankton functional groups is one of the key scientific issues. Therefore, it is critical to study the composition and ecological characteristics of phytoplankton functional groups (PFGs) in the Yellow Sea (YS) and the East China Sea (ECS).
From April 2006 to June 2008, nine cruises were conducted in the YS and the ECS. Based on phytoplankton data by Unterm?hl method, composition and ecological characteristics of PFGs were investigated. Relationships between PFGs and environmental factors were analyzed by CCA, and influences of hydrodynamic processes on PFGs were taken in account as well.
According to trophic function, phytoplankton is divided into 12 functional groups, i.e. pico-phytoplankton (PicoPl), nano-diatom (NanDiat), autotrophic nano-flagellate (NanAutFl), autotrophic micro-dinoflagellate (MAutFl), mixotrophic micro-dinoflagellate (MMixFl), heterotrophic micro-dinoflagellate (MHetFl), unicellular micro-diatom (MUniDiat), hair-shaped micro-diatom (MHairDiat), branch-shaped micro-diatom (MBranDiat), toxic phytoplankton (ToxPl), inedible phytoplankton (InedPl), and other phytoplankton (Others).
Phytoplankton integrated biomass in spring was much higher than that in autumn in the southern YS. High biomass distributed in the coastal water of Shandong Peninsula in spring, whereas, in the coastal water of Cheju Island in autumn. Phytoplankton integrated biomass showed evident seasonal variation in the Yangtze River Estuary (YRE) and its adjacent waters, i.e., in summer > in spring > in autumn > in winter. Biomass in the coastal waters of YRE and Zhejiang Province was much higher than that in off-shore sites with salinity > 31. Vertically, phytoplankton biomass decreased with depth from surface to bottom.
Composition of PFGs varied spatially and temporally in the YS and the ECS. MHairDiat was the most dominant group in the southern YS in spring and autumn. MHetFl-dominant dinoflagellate groups were most important, in addition, ToxPl was also important in the YRE and its adjacent waters in late spring. Diatom groups replaced dinoflagellate, and MHairDiat and MBranDiat were dominant in the YRE and its adjacent waters in the middle of summer. Diatom groups accounted for 95% of phytoplankton biomass in the YRE and its adjacent waters in late summer. Diatom groups, MHairDiat > MUniDiat > MBranDiat, were dominant in the continental shelf water of ECS in autumn and winter. Dinoflagellate groups, MMixFl > MAutFl > MHetFl, were dominant in the YRE and the coastal ECS in May of 2007, while MHetFl > MMixFl > MAutFl in June of 2008.
MHairDiat was most abundant in the YS Trough water near Shandong Peninsula in spring. Its biomass in the YRE and the coastal ECS increased from late spring to late summer, and decreased from autumn to winter. MHairDiat showed extremely low biomass in the Yellow Sea Cold Water Mass (YSCWM) area in autumn. Distribution of MUniDiat was similar to that of MHairDiat in spring. Its biomass in the YRE and the coastal ECS increased from August to October. MUniDiat was more abundant in the ECS than in the southern YS in autumn. MUniDiat biomass in winter was lower than that in autumn in the ECS, and was densely distributed near the YRE. In spring, MBranDiat was abundant in the YS water near the YRE, while scarce in the ECS. MBranDiat biomass increased from June to October, and decreased from autumn to winter in the ECS. Late spring peak value appeared in coastal water of Hangzhou Bay, while autumn peak value in the open sea. In autumn MBranDiat was densely distributed in the coastal water of YS. High density of MHetFl located in coastal water of Shandong Peninsula and the area near the YRE and Hangzhou Bay in spring. MHetFl biomass was much low in the YS and the ECS from summer to winter. MMixFl biomass was highest in late spring among four seasons, and its peak value occurred in the YRE and the coastal ECS. MMixFl biomass was higher in the ECS than in the YS in spring, while lower in the ECS than in the YS in autumn. MMixFl biomass was extremely low in the continental shelf water of ECS in winter. Maximum MAutFl biomass occurred in the YRE adjacent YS in late spring, and high biomass decreased in summer. MAutFl was in low abundance in the continental shelf water of ECS in autumn and winter. ToxPl was abundant in the middle of southern YS, in the YRE and the coastal ECS in spring, and showed peak biomass, which disappeared in late summer, in the YRE adjacent YS in the middle of summer. InedPl was most abundant in the YRE adjacent YS in late summer. Generally, its biomass was high in open sea.
Succession of keystone species was evident in survey area. Keystone species were mainly diatoms in the southern YS. Keystone species were Thalassiosira pacifica and Corethrom hystrix in spring, and Guinardia striata and Coscinodiscus sp. in autumn. In the YRE and the ECS, dinoflagellates, such as Noctiluca scientillans, Prorocentrum dentatum, Karenia mikimotoi, became keystone species in May and June, besides toxic phytoplankton Pseudo-nitzschia delicatissima showed high biomass as well. Diatom, i.e. Dactyliosolen fragilissimus, Chaetoceros curvisetus, Rhizosolenia crassipina, Lauderia annulata, replaced dinoflagellate and became keystone species in summer. Diatoms, such as Coscinodiscus sp., Thalassiosira rotula, Guinardia striata, became keystone species again in autumn. Keystone species in winter was mainly unicellular diatom, such as Coscinodiscus sp.. Noctiluca scientillans, Prorocentrum dentatum, and Karenia mikimotoi could bloom in the YRE and its adjacent water in spring, so could Thalassiosira pacifica in the southern YS in spring.
Correlations between PFGs and environmental factors were highly variable in spatial and temporal scale. MHairDiat was well correlated with salinity and temperature in survey area, however, it could be limited by nutrients during cruise LST-I and cruise QYQ-II. Distributions of MUniDiat were mainly influenced by nutrients, but also correlated with salinity during cruise LST-I. Distributions of MBranDiat could be influenced by nutrients in spring and summer, while by temperature and salinity in late summer, autumn and winter. Phosphate was the key factor that influenced the distributions of MAutFl during cruise LST-I, while ammonia in the Yangtze River Estuary, and nutrients during cruise ZLJ-I, and temperature and salinity during other cruises. MMixFl well correlated with temperature and salinity, whereas correlated with nitrogen during cruise LST-I and cruise ZLJ-I. MHetFl mainly distributed in upper layers of coastal waters, however, it could show high biomass in open sea as well. ToxPl was mainly influenced by nitrogen during cruise LST-I, cruise QYQ-II, and cruise ZLJ-II. Distributions of InedPl were mostly controlled by temperature and salinity, in addition, InedPl positively correlated with nutrient during cruise ZLJ-I and cruise ZLJ-II.
Phytoplankton biomass was extremely low in the YSCWM in autumn, where the strong thermocline stopped the upward transport of nutrients from bottom water. There was mainly phosphate limitation in surface water, where MMixFl and MHetFl were dominant, and illumination limitation below surface, where MUniDiat was dominant. High phytoplankton biomass in the YRE was supported by rich nutrients transported by Yangtze River Dilute Water. Phytoplankton biomass front appeared in the waters where nutrients and illumination stroke a balance. Phytoplankton biomass was generally low in Kuroshio and Taiwan Warm Current, characteristic of high salinity and low nutrients.
Cluster analysis of survey sites of every cruise was conducted based on PFGs composition. According to cluster analysis and phytoplankton biomass, the YS and the ECS were divieded into several provinces. The southern YS were divided into 4 provinces in spring, while 3 provinces in autumn. The YRE and its adjacent YS and ECS was divided into 2 provinces in late spring, and 4 provinces in the middle of summer, and 3 provinces in late summer. The continental shelf water of ECS was divided into 2 provinces in both autumn and winter. The YRE and the coastal ECS were divided into 2 provinces in spring and 3 provinces in late spring.
PFGs provinces were similar to zooplankton functional groups provinces in the southern YS in spring and autumn, however, they were quiet different in distribution of biomass. Biomass and composition of PFGs was closely correlated with grazing, metabolism, and reproduction of Calanus sinicus. High biomass of MHairDiat and MBranDiat could be a key factor in formation of hypoxia in the YRE in summer.
Based on carbon biomass, the composition and distributions of PFGs were reported in the YS and the ECS, so was the distribution of keystone species. Correlations between PFGs and environmental factors were analyzed, and PFGs provinces were divided as well. This dissertation provides for the food output model in the YS and the ECS. To understand the PFGs comprehensively, multi-method should be employed in the analysis of phytoplankton samples. Furthermore, interdisciplinarity study should be conducted to get insights into trophic structure of pelagic food web in the YS and the ECS.
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