我国海区浮游纤毛虫的生态学研究
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摘要
海洋浮游纤毛虫是在海洋中营浮游生活的纤毛虫,主要是舞毛亚纲(Choreotrichia)和寡毛亚纲(Oligotrichia)的种类。浮游纤毛虫的粒级范围为10-200μm,平均丰度约为102-103 ind. L-1,广泛分布于各种生境中。浮游纤毛虫是海洋浮游生态系统中小型浮游动物的主要类群之一,它不仅是pico-、nano-级浮游生物等初级生产者的主要摄食者,而且是中型浮游动物(如桡足类)的饵料,是微食物网向经典食物链物质和能量传递的关键环节。
我国海区浮游纤毛虫生态学的基础资料还较少,许多海区如南海还缺乏浮游纤毛虫丰度和生物量的基础资料,东海已有的研究比较零散,黄海春季水华过程还没有纤毛虫丰度和生物量时空变化的报道。此外,关于纤毛虫的摄食作用的研究更少。本文报道了南海和东海纤毛虫丰度和生物量以及分布;探讨了东海水团对纤毛虫分布的影响;研究了纤毛虫在春季水华过程中的时空变化;并在黄海进行了小型浮游动物对浮游植物和鞭毛虫的摄食,桡足类(中华哲水蚤)对纤毛虫的摄食研究。
不同海区纤毛虫的丰度和生物量
2007年10月南海北部海域纤毛虫丰度为0-5757 (848±776) ind. L-1,无壳纤毛虫占绝对优势,其丰度占总丰度的比例平均为91.9±9%。纤毛虫生物量为0-12.09 (1.2μ1.54)μg C L-1,无壳纤毛虫的生物量平均为0.94±1.27μg C L-1,占总生物量的78.6μ23.8%。共发现砂壳纤毛虫16个属,49种,拟铃虫属的种类最多。纤毛虫多分布于近岸浅水区(高温低盐,高Chl a浓度)。纤毛虫最大丰度5757 ind. L-1高于我国其他海区的调查研究。
在长江口及毗邻海域,2006年8月纤毛虫丰度为0-4163 (718±571) ind. L-1,生物量为0.00-64.88 (1.86±6.34)μg C L-1;10月纤毛虫丰度为35-1155 (358±235) ind. L-1,生物量为0.00-9.23 (0.5±1.15)μg C L-1。8月表层和水柱纤毛虫丰度、生物量高值出现在长江口和杭州湾口以东近岸,10月高值出现在远岸,且南部高于北部。10月砂壳纤毛虫占总纤毛虫生物量的比例略高于8月。DO (溶解氧,Dissolved Oxygen)浓度对纤毛虫水体垂直分布的影响不明显。
在东海陆架区,2006年11-12月(秋季)纤毛虫丰度为0-1795 (208±266) ind. L-1,生物量为0-2.36 (0.28±0.35)μg C L-1;2007年2-3月(冬季)纤毛虫丰度为0-22695 (524±1990) ind. L-1,生物量为0-10.87 (0.47±1.01)μg C L-1。纤毛虫丰度与生物量秋季在外陆架区和中陆架区高于内陆架区,冬季中陆架区高于外陆架区和内陆架区。两个季节都是无壳纤毛虫的丰度占优势,而秋季砂壳纤毛虫对生物量的贡献大于无壳纤毛虫。ESD (Equivalent Spherical Diameter) 10-20μm的小型纤毛虫分别占秋季和冬季纤毛虫丰度的63%和82%。与20世纪90年代的调查研究相比,东海陆架区浮游纤毛虫的生态分布没有发生明显变化。
纤毛虫丰度和水团的关系
纤毛虫丰度在东海的分布受到水团的影响。2006年8月(夏季)在长江口及毗邻海域,冲淡水表层纤毛虫平均丰度(972±746 ind. L-1)高于远岸的混合水(475±345 ind. L-1)。盐度是影响混合水纤毛虫表层丰度分布的重要因素。2007年2-3月(冬季)在陆架区,混合水的表层纤毛虫平均丰度(452±645 ind. L-1)高于沿岸水(209±307 ind. L-1)和黑潮水(202±170 ind. L-1)。混合水的纤毛虫0-30 m水柱丰度与盐度呈显著的负相关,与Chl a浓度呈显著的正相关。纤毛虫丰度在锋面区附近比较高。夏季长江口及毗邻海域发现的砂壳纤毛虫大多为近岸浅水种,其分布在两个水团没有明显的分区;而冬季在陆架区砂壳纤毛虫的分布呈现明显的分区,有些偶见种可能指示了黑潮水入侵陆架的路径。
黄海春季水华过程中纤毛虫水柱生物量的变化
在不同类型的水华过程纤毛虫和红色中缢虫的丰度和生物量有显著不同。2006年在典型硅藻水华站,纤毛虫水柱生物量高于2007年和2009年水华过程中的最大水柱生物量。
2007年,发现三次硅藻的水华过程(BM1站、BM2站和BM3站),在BM1站,红色中缢虫丰度达2.9±105 ind. L-1。在BM1站和BM2站水华过程中,红色中缢虫丰度和水柱生物量都迅速降低到低值,纤毛虫(不包括中缢虫)平均水柱生物量也降低。BM1和BM2水华过程的红色中缢虫平均水柱生物量高于纤毛虫;在BM3站水华过程中红色中缢虫丰度很低,其水柱生物量低于纤毛虫。
2009年,在硅藻水华过程中,眼虫最大丰度3.5×104 ind. L-1,红色中缢虫丰度较低,纤毛虫平均水柱生物量远高于红色中缢虫,在水华的后期有明显的增加;而在甲藻和硅藻的混合水华过程中,红色中缢虫平均水柱生物量高于纤毛虫,纤毛虫和红色中缢虫的日平均水柱生物量都在第四天达到最大值然后降低。
小型浮游动物对浮游植物和鞭毛虫的摄食
2007年4月,在水华BM1-1站,浮游植物的生长率(1.18 d-1)和小型浮游动物的摄食死亡率(0.76 d-1)都高于水华后期和非水华站,小型浮游动物对初级生产力的摄食压力低于水华后期和多数的非水华站。纤毛虫和环沟藻在水华过程中的平均水柱生物量不高于非水华站。因此,小型浮游动物对春季水华的下行控制作用可能不明显。
2007年5月在南黄海,稀释培养后纤毛虫丰度与稀释因子呈线性关系;而鞭毛虫丰度在稀释因子大时增加。在富营养站,小型浮游动物对鞭毛虫的摄食率(1.01 dμ1)高于寡营养站(0.42-0.44 dμ1)。在富营养站位,绝大多数的鞭毛虫生产力(99%)可以被小型浮游动物摄食,而在寡营养站位,小型浮游动物摄食72%-73%的鞭毛虫生产力。
桡足类对纤毛虫的摄食
2009年4月,在硅藻水华过程中,水华中期(B20-29站)中华哲水蚤对纤毛虫和红色中缢虫的清滤率(155 ml ind.-1 d-1,252 ml ind.-1 d-1)分别高于水华后期(B20-85站,20 ml ind.-1 d-1,147 ml ind.-1 d-1)和非水华期(B31站,98 ml ind.-1 d-1,79 ml ind.-1 d-1)。水华中期中华哲水蚤对纤毛虫和红色中缢虫现存量的摄食压力(83.3%,94.6%)也分别高于非水华B31站(67.8%, 59.8%)和水华后期(20.9%, 81.7%)。桡足类摄食可能是影响水华过程中纤毛虫生物量变化的重要因素。
2007年5月,在南黄海寡营养站,桡足类添加培养实验中中华哲水蚤的添加明显降低了纤毛虫的丰度,而鞭毛虫的丰度没有相应地增加,蓝细菌丰度明显增加。因此,在桡足类-纤毛虫-鞭毛虫食物链中没有发现营养级联效应,纤毛虫可能是蓝细菌的主要摄食者。
Marine planktonic ciliates mainly include subclass Choreotrichia and Oligotrichia. They are unicellular and eukaryotic protists in the size range of 10-200μm. They are abundant and ubiquitous with average abundance 102-103 ind. L-1 in various marine habitats. As one of the major components of microzooplankton in the pelagic ecosystem, ciliates play a key role in the transfer of carbon and energy through microbial food web to classical food chain, acting as top-down consumer of, pico-, nano- plankton and other primary producers and as a food source for mesozooplankton (e.g. copepod).
Ecological researches of planktonic ciliate in the coastal area of China seas are limited. Basic data of ciliate abundance and biomass is scarce in the South China Sea; those in the East China Sea are scattered. There is no research on the spatio-temporal variability of ciliate abundance and biomass during blooms and much less reports on grazing impact of ciliate. In this study, abundance, biomass and distribution of planktonic ciliate were investigated in the South China Sea and East China Sea. Influence of different water masses on distribution of ciliate abundance was studied in the East China Sea. Spatio-temporal variability of ciliate abundance and biomass during spring blooms was investigated in the Yellow Sea. In addition, incubations were carried out to study the grazing of microzooplankton on phytoplankton and nanoflagellate, the grazing of copepod (Calanus sinicus) on ciliate.
Ciliate abundance and biomass in different seas
In the northern South China Sea in October, 2007, ciliate abundance ranged from 0 to 5757 (on average 848±776) ind. L-1, of which dominant aloricate ciliates occupied 91.9±9%. The biomass of all ciliates was 0-2.09 (on average 1.2±1.54)μg C L-1, of which aloricate ciliates (on average 0.94±1.27μg C L-1) occupied 78.6μ23.8% to the total. Forty-nine species of tintinnids in 16 genera were identified. Genus Tintinnopsis was dominant in abundance. Ciliates mainly distributed near coastal shallow waters which was warm and less salty with more Chl a concentration. The maximum abundance of ciliate in the study was higher than those in other parts of China seas.
In the Changjiang River Estuary and its adjacent sea, ciliate abundance ranged from 0 to 4163 (on average 718±571) ind. L-1 and the biomass ranged from 0.00 to 64.88 (on average 1.86±6.34)μg C L-1 in August 2006; the ciliate abundance ranged from 35 to 1155 (on average 358±235) ind. L-1 and the biomass ranged from 0.00 to 9.23 (on average 0.5±1.15)μg C L-1 in October 2006. Higher surface and integrated abundance and biomass of ciliate occurred in the coastal water in the eastern of Changjiang River mouth and Hangzhou Bay mouth in August. However, those occurring in the offshore in October were higher in the northern than in the southern part. The contribution of tintinnid to total biomass in October was slightly higher than in August. Influence of dissolved oxygen (DO) concentration on the vertical distribution of ciliate abundance and biomass was not significant.
In the shelf areas of East China Sea, ciliate abundance and biomass ranged from 0 to 1795 (on average 208μ266) ind. L-1 and from 0 to 2.36 (on average 0.28μ0.35)μg C L-1, respectively, in autumn (11.19-12.23), 2006. Ciliate abundance and biomass were in the range of 0 to 22695 (on average 524μ1990) ind. L-1 and 0 to 10.87 (on average 0.47μ1.01)μg C L-1, respectively, in winter (2.22-3.11), 2007. In autumn, ciliate abundance and biomass in the outer and middle shelf were higher than in the inner shelf. However, in winter, more ciliates occurred in the middle shelf than in the outer and inner shelf. Aloricate ciliates were dominant in the abundance in both autumn and winter, but contribution of tintinnid to total biomass was higher than aloricate ciliate in autumn. Small ciliates (ESD 10-20μm) accounted for 63% and 82% in abundance in autumn and winter, respectively. In comparison with reports in the 1990s in East China Sea, no distinct variation on ecological distribution of planktonic ciliates was found.
Distribution of ciliate abundance in relation to water masses
In the East China Sea, distribution of ciliate abundance was affected by different water masses. In August (summer) 2006, average surface ciliate abundance (972±746 ind. L-1) in the Changjiang Diluted Water (CDW) was higher than in the offshore Shelf Mixing Water (SMW) (475±345 ind. L-1) in the Changjiang River Estuary and its adjacent sea. Salinity variation had important effects on the spatial pattern of surface ciliate abundance in the SMW.
In February-March (winter) 2007, average surface ciliate abundance in the SMW (452μ645 ind. L-1) was higher than in the Kuroshio Water (202±170 ind. L-1) and the Coastal Water (CoW) (209±307 ind. L-1) in the shelf area. Integrated ciliate abundance of 0-30 m in the SMW was not only significantly and negatively correlated with salinity, but also positively correlated with Chl a concentration.
Ciliate distribution was characterized by increase of abundance close to the frontal areas. Most tintinnids identified as neritic species did not show discrimination of distribution between two water masses in the Changjiang River Estuary in summer. However, there were pronounced distribution zones of tintinnid species and some occasional species might indicate the intrusion route of Kuroshio Water on the continental shelf in winter.
Variability of ciliate integrated biomass during spring blooms in the Yellow Sea
Ciliate and Myrionecta rubra abundance and biomass responded differently to various phytoplankton blooms.
Ciliate integrated biomass at representative diatom bloom station in 2006 was higher than the maximum integrated biomass during blooms in 2007 and 2009.
In 2007, diatom blooms at three stations (St. BM1, St. BM2 and St. BM3) were found. At St. BM1, maximum abundance of M. rubra was up to 2.9×105 ind. L-1. During the bloom of St. BM1 and St. BM2, M. rubra abundance and integrated biomass sharply decreased. Non-Myrionecta ciliates daily average integrated biomass decreased. Average integrated biomass of M. rubra was higher than ciliate during the bloom of St. BM1 and St. BM2, but during the bloom of St. BM3, integrated biomass of M. rubra with low abundance was lower than ciliate.
In 2009, M. rubra abundance was low during the diatom bloom with Euglena sp. occurring in the maximum abundance 3.5×104 ind. L-1. Average integrated biomass of ciliate that was much higher than M. rubra increased at the end of bloom. During the dinoflagellate and diatom mixed bloom, average integrated biomass of M. rubra was higher than ciliate. Daily average integrated biomass of ciliate and M. rubra reached a peak on the fourth day and then declined.
Grazing of microzooplankton on phytoplankton and nanoflagellate
In April 2007, phytoplankton growth rate (1.18 d-1) and microzooplankton grazing mortality rate (0.76 d-1) at bloom St. BM1-1 were higher than those at the post-bloom and non-bloom stations. Microzooplankton daily grazing impact on primary production at St. BM1-1 was lower than post-bloom and most of non-bloom stations. Average integrated biomass of ciliate and Gyrodinium sp. during the bloom were not higher than those at non-bloom stations. Therefore, the top-down control of microzooplankton on spring bloom might be not significant.
In May 2007,ciliate abundance kept a proportional relationship with dilution factors after incubation in the dilution incubations while flagellate abundance increased in the more diluted series at both eutrophic and oligotrophic stations in the southern Yellow Sea. Microzooplankton grazing rate on flagellates was higher in the eutrophic water (1.01 d~（-1）) than in the oligotrophic water (0.42-0.44 d~（-1）). In the eutrophic water, most of the flagellate production (99%) was grazed by microzooplankton. In the oligotrophic water, only 72-73% of the flagellate production was grazed by microzooplankton. Grazing of Copepod on ciliate
In April 2009, clearance rates of ciliates and M. rubra by C. sinicus (155 ml copepod~（-1）d~（-1）, 252 ml copepod~（-1）d~（-1）) during the course of bloom (St. B20-29) were higher than those at the end of bloom (St. B20-85, 20 ml ind.~（-1）d~（-1）, 147 ml ind.~（-1）d~（-1）) and at non-bloom stations (St. B31, 98 ml ind.~（-1）d~（-1）, 79 ml ind.~（-1）d~（-1）), respectively. Daily grazing pressures on ciliate (83.3%) and M. rubra (94.6%) during the bloom (St. B20-29) were higher than those at non-bloom station (67.8%, 59.8%) and at the end of bloom (20.9%, 81.7%), respectively. Copepod grazing may be an important factor in the variability of ciliate biomass during the bloom.
In May 2007, C. sinicus additions in the incubation significantly reduced ciliate abundance at the oligotrophic stations in the the southern Yellow Sea. Heterotrophic flagellates did not increase while Synechococcus abundance.increased significantly. Therefore, there was no trophic cascade along the copepod-ciliate-flagellate food chain. Ciliates might be the dominant grazers of Synechococcus.
我国海区浮游纤毛虫生态学的基础资料还较少,许多海区如南海还缺乏浮游纤毛虫丰度和生物量的基础资料,东海已有的研究比较零散,黄海春季水华过程还没有纤毛虫丰度和生物量时空变化的报道。此外,关于纤毛虫的摄食作用的研究更少。本文报道了南海和东海纤毛虫丰度和生物量以及分布;探讨了东海水团对纤毛虫分布的影响;研究了纤毛虫在春季水华过程中的时空变化;并在黄海进行了小型浮游动物对浮游植物和鞭毛虫的摄食,桡足类(中华哲水蚤)对纤毛虫的摄食研究。
不同海区纤毛虫的丰度和生物量
2007年10月南海北部海域纤毛虫丰度为0-5757 (848±776) ind. L-1,无壳纤毛虫占绝对优势,其丰度占总丰度的比例平均为91.9±9%。纤毛虫生物量为0-12.09 (1.2μ1.54)μg C L-1,无壳纤毛虫的生物量平均为0.94±1.27μg C L-1,占总生物量的78.6μ23.8%。共发现砂壳纤毛虫16个属,49种,拟铃虫属的种类最多。纤毛虫多分布于近岸浅水区(高温低盐,高Chl a浓度)。纤毛虫最大丰度5757 ind. L-1高于我国其他海区的调查研究。
在长江口及毗邻海域,2006年8月纤毛虫丰度为0-4163 (718±571) ind. L-1,生物量为0.00-64.88 (1.86±6.34)μg C L-1;10月纤毛虫丰度为35-1155 (358±235) ind. L-1,生物量为0.00-9.23 (0.5±1.15)μg C L-1。8月表层和水柱纤毛虫丰度、生物量高值出现在长江口和杭州湾口以东近岸,10月高值出现在远岸,且南部高于北部。10月砂壳纤毛虫占总纤毛虫生物量的比例略高于8月。DO (溶解氧,Dissolved Oxygen)浓度对纤毛虫水体垂直分布的影响不明显。
在东海陆架区,2006年11-12月(秋季)纤毛虫丰度为0-1795 (208±266) ind. L-1,生物量为0-2.36 (0.28±0.35)μg C L-1;2007年2-3月(冬季)纤毛虫丰度为0-22695 (524±1990) ind. L-1,生物量为0-10.87 (0.47±1.01)μg C L-1。纤毛虫丰度与生物量秋季在外陆架区和中陆架区高于内陆架区,冬季中陆架区高于外陆架区和内陆架区。两个季节都是无壳纤毛虫的丰度占优势,而秋季砂壳纤毛虫对生物量的贡献大于无壳纤毛虫。ESD (Equivalent Spherical Diameter) 10-20μm的小型纤毛虫分别占秋季和冬季纤毛虫丰度的63%和82%。与20世纪90年代的调查研究相比,东海陆架区浮游纤毛虫的生态分布没有发生明显变化。
纤毛虫丰度和水团的关系
纤毛虫丰度在东海的分布受到水团的影响。2006年8月(夏季)在长江口及毗邻海域,冲淡水表层纤毛虫平均丰度(972±746 ind. L-1)高于远岸的混合水(475±345 ind. L-1)。盐度是影响混合水纤毛虫表层丰度分布的重要因素。2007年2-3月(冬季)在陆架区,混合水的表层纤毛虫平均丰度(452±645 ind. L-1)高于沿岸水(209±307 ind. L-1)和黑潮水(202±170 ind. L-1)。混合水的纤毛虫0-30 m水柱丰度与盐度呈显著的负相关,与Chl a浓度呈显著的正相关。纤毛虫丰度在锋面区附近比较高。夏季长江口及毗邻海域发现的砂壳纤毛虫大多为近岸浅水种,其分布在两个水团没有明显的分区;而冬季在陆架区砂壳纤毛虫的分布呈现明显的分区,有些偶见种可能指示了黑潮水入侵陆架的路径。
黄海春季水华过程中纤毛虫水柱生物量的变化
在不同类型的水华过程纤毛虫和红色中缢虫的丰度和生物量有显著不同。2006年在典型硅藻水华站,纤毛虫水柱生物量高于2007年和2009年水华过程中的最大水柱生物量。
2007年,发现三次硅藻的水华过程(BM1站、BM2站和BM3站),在BM1站,红色中缢虫丰度达2.9±105 ind. L-1。在BM1站和BM2站水华过程中,红色中缢虫丰度和水柱生物量都迅速降低到低值,纤毛虫(不包括中缢虫)平均水柱生物量也降低。BM1和BM2水华过程的红色中缢虫平均水柱生物量高于纤毛虫;在BM3站水华过程中红色中缢虫丰度很低,其水柱生物量低于纤毛虫。
2009年,在硅藻水华过程中,眼虫最大丰度3.5×104 ind. L-1,红色中缢虫丰度较低,纤毛虫平均水柱生物量远高于红色中缢虫,在水华的后期有明显的增加;而在甲藻和硅藻的混合水华过程中,红色中缢虫平均水柱生物量高于纤毛虫,纤毛虫和红色中缢虫的日平均水柱生物量都在第四天达到最大值然后降低。
小型浮游动物对浮游植物和鞭毛虫的摄食
2007年4月,在水华BM1-1站,浮游植物的生长率(1.18 d-1)和小型浮游动物的摄食死亡率(0.76 d-1)都高于水华后期和非水华站,小型浮游动物对初级生产力的摄食压力低于水华后期和多数的非水华站。纤毛虫和环沟藻在水华过程中的平均水柱生物量不高于非水华站。因此,小型浮游动物对春季水华的下行控制作用可能不明显。
2007年5月在南黄海,稀释培养后纤毛虫丰度与稀释因子呈线性关系;而鞭毛虫丰度在稀释因子大时增加。在富营养站,小型浮游动物对鞭毛虫的摄食率(1.01 dμ1)高于寡营养站(0.42-0.44 dμ1)。在富营养站位,绝大多数的鞭毛虫生产力(99%)可以被小型浮游动物摄食,而在寡营养站位,小型浮游动物摄食72%-73%的鞭毛虫生产力。
桡足类对纤毛虫的摄食
2009年4月,在硅藻水华过程中,水华中期(B20-29站)中华哲水蚤对纤毛虫和红色中缢虫的清滤率(155 ml ind.-1 d-1,252 ml ind.-1 d-1)分别高于水华后期(B20-85站,20 ml ind.-1 d-1,147 ml ind.-1 d-1)和非水华期(B31站,98 ml ind.-1 d-1,79 ml ind.-1 d-1)。水华中期中华哲水蚤对纤毛虫和红色中缢虫现存量的摄食压力(83.3%,94.6%)也分别高于非水华B31站(67.8%, 59.8%)和水华后期(20.9%, 81.7%)。桡足类摄食可能是影响水华过程中纤毛虫生物量变化的重要因素。
2007年5月,在南黄海寡营养站,桡足类添加培养实验中中华哲水蚤的添加明显降低了纤毛虫的丰度,而鞭毛虫的丰度没有相应地增加,蓝细菌丰度明显增加。因此,在桡足类-纤毛虫-鞭毛虫食物链中没有发现营养级联效应,纤毛虫可能是蓝细菌的主要摄食者。
Marine planktonic ciliates mainly include subclass Choreotrichia and Oligotrichia. They are unicellular and eukaryotic protists in the size range of 10-200μm. They are abundant and ubiquitous with average abundance 102-103 ind. L-1 in various marine habitats. As one of the major components of microzooplankton in the pelagic ecosystem, ciliates play a key role in the transfer of carbon and energy through microbial food web to classical food chain, acting as top-down consumer of, pico-, nano- plankton and other primary producers and as a food source for mesozooplankton (e.g. copepod).
Ecological researches of planktonic ciliate in the coastal area of China seas are limited. Basic data of ciliate abundance and biomass is scarce in the South China Sea; those in the East China Sea are scattered. There is no research on the spatio-temporal variability of ciliate abundance and biomass during blooms and much less reports on grazing impact of ciliate. In this study, abundance, biomass and distribution of planktonic ciliate were investigated in the South China Sea and East China Sea. Influence of different water masses on distribution of ciliate abundance was studied in the East China Sea. Spatio-temporal variability of ciliate abundance and biomass during spring blooms was investigated in the Yellow Sea. In addition, incubations were carried out to study the grazing of microzooplankton on phytoplankton and nanoflagellate, the grazing of copepod (Calanus sinicus) on ciliate.
Ciliate abundance and biomass in different seas
In the northern South China Sea in October, 2007, ciliate abundance ranged from 0 to 5757 (on average 848±776) ind. L-1, of which dominant aloricate ciliates occupied 91.9±9%. The biomass of all ciliates was 0-2.09 (on average 1.2±1.54)μg C L-1, of which aloricate ciliates (on average 0.94±1.27μg C L-1) occupied 78.6μ23.8% to the total. Forty-nine species of tintinnids in 16 genera were identified. Genus Tintinnopsis was dominant in abundance. Ciliates mainly distributed near coastal shallow waters which was warm and less salty with more Chl a concentration. The maximum abundance of ciliate in the study was higher than those in other parts of China seas.
In the Changjiang River Estuary and its adjacent sea, ciliate abundance ranged from 0 to 4163 (on average 718±571) ind. L-1 and the biomass ranged from 0.00 to 64.88 (on average 1.86±6.34)μg C L-1 in August 2006; the ciliate abundance ranged from 35 to 1155 (on average 358±235) ind. L-1 and the biomass ranged from 0.00 to 9.23 (on average 0.5±1.15)μg C L-1 in October 2006. Higher surface and integrated abundance and biomass of ciliate occurred in the coastal water in the eastern of Changjiang River mouth and Hangzhou Bay mouth in August. However, those occurring in the offshore in October were higher in the northern than in the southern part. The contribution of tintinnid to total biomass in October was slightly higher than in August. Influence of dissolved oxygen (DO) concentration on the vertical distribution of ciliate abundance and biomass was not significant.
In the shelf areas of East China Sea, ciliate abundance and biomass ranged from 0 to 1795 (on average 208μ266) ind. L-1 and from 0 to 2.36 (on average 0.28μ0.35)μg C L-1, respectively, in autumn (11.19-12.23), 2006. Ciliate abundance and biomass were in the range of 0 to 22695 (on average 524μ1990) ind. L-1 and 0 to 10.87 (on average 0.47μ1.01)μg C L-1, respectively, in winter (2.22-3.11), 2007. In autumn, ciliate abundance and biomass in the outer and middle shelf were higher than in the inner shelf. However, in winter, more ciliates occurred in the middle shelf than in the outer and inner shelf. Aloricate ciliates were dominant in the abundance in both autumn and winter, but contribution of tintinnid to total biomass was higher than aloricate ciliate in autumn. Small ciliates (ESD 10-20μm) accounted for 63% and 82% in abundance in autumn and winter, respectively. In comparison with reports in the 1990s in East China Sea, no distinct variation on ecological distribution of planktonic ciliates was found.
Distribution of ciliate abundance in relation to water masses
In the East China Sea, distribution of ciliate abundance was affected by different water masses. In August (summer) 2006, average surface ciliate abundance (972±746 ind. L-1) in the Changjiang Diluted Water (CDW) was higher than in the offshore Shelf Mixing Water (SMW) (475±345 ind. L-1) in the Changjiang River Estuary and its adjacent sea. Salinity variation had important effects on the spatial pattern of surface ciliate abundance in the SMW.
In February-March (winter) 2007, average surface ciliate abundance in the SMW (452μ645 ind. L-1) was higher than in the Kuroshio Water (202±170 ind. L-1) and the Coastal Water (CoW) (209±307 ind. L-1) in the shelf area. Integrated ciliate abundance of 0-30 m in the SMW was not only significantly and negatively correlated with salinity, but also positively correlated with Chl a concentration.
Ciliate distribution was characterized by increase of abundance close to the frontal areas. Most tintinnids identified as neritic species did not show discrimination of distribution between two water masses in the Changjiang River Estuary in summer. However, there were pronounced distribution zones of tintinnid species and some occasional species might indicate the intrusion route of Kuroshio Water on the continental shelf in winter.
Variability of ciliate integrated biomass during spring blooms in the Yellow Sea
Ciliate and Myrionecta rubra abundance and biomass responded differently to various phytoplankton blooms.
Ciliate integrated biomass at representative diatom bloom station in 2006 was higher than the maximum integrated biomass during blooms in 2007 and 2009.
In 2007, diatom blooms at three stations (St. BM1, St. BM2 and St. BM3) were found. At St. BM1, maximum abundance of M. rubra was up to 2.9×105 ind. L-1. During the bloom of St. BM1 and St. BM2, M. rubra abundance and integrated biomass sharply decreased. Non-Myrionecta ciliates daily average integrated biomass decreased. Average integrated biomass of M. rubra was higher than ciliate during the bloom of St. BM1 and St. BM2, but during the bloom of St. BM3, integrated biomass of M. rubra with low abundance was lower than ciliate.
In 2009, M. rubra abundance was low during the diatom bloom with Euglena sp. occurring in the maximum abundance 3.5×104 ind. L-1. Average integrated biomass of ciliate that was much higher than M. rubra increased at the end of bloom. During the dinoflagellate and diatom mixed bloom, average integrated biomass of M. rubra was higher than ciliate. Daily average integrated biomass of ciliate and M. rubra reached a peak on the fourth day and then declined.
Grazing of microzooplankton on phytoplankton and nanoflagellate
In April 2007, phytoplankton growth rate (1.18 d-1) and microzooplankton grazing mortality rate (0.76 d-1) at bloom St. BM1-1 were higher than those at the post-bloom and non-bloom stations. Microzooplankton daily grazing impact on primary production at St. BM1-1 was lower than post-bloom and most of non-bloom stations. Average integrated biomass of ciliate and Gyrodinium sp. during the bloom were not higher than those at non-bloom stations. Therefore, the top-down control of microzooplankton on spring bloom might be not significant.
In May 2007,ciliate abundance kept a proportional relationship with dilution factors after incubation in the dilution incubations while flagellate abundance increased in the more diluted series at both eutrophic and oligotrophic stations in the southern Yellow Sea. Microzooplankton grazing rate on flagellates was higher in the eutrophic water (1.01 d~（-1）) than in the oligotrophic water (0.42-0.44 d~（-1）). In the eutrophic water, most of the flagellate production (99%) was grazed by microzooplankton. In the oligotrophic water, only 72-73% of the flagellate production was grazed by microzooplankton. Grazing of Copepod on ciliate
In April 2009, clearance rates of ciliates and M. rubra by C. sinicus (155 ml copepod~（-1）d~（-1）, 252 ml copepod~（-1）d~（-1）) during the course of bloom (St. B20-29) were higher than those at the end of bloom (St. B20-85, 20 ml ind.~（-1）d~（-1）, 147 ml ind.~（-1）d~（-1）) and at non-bloom stations (St. B31, 98 ml ind.~（-1）d~（-1）, 79 ml ind.~（-1）d~（-1）), respectively. Daily grazing pressures on ciliate (83.3%) and M. rubra (94.6%) during the bloom (St. B20-29) were higher than those at non-bloom station (67.8%, 59.8%) and at the end of bloom (20.9%, 81.7%), respectively. Copepod grazing may be an important factor in the variability of ciliate biomass during the bloom.
In May 2007, C. sinicus additions in the incubation significantly reduced ciliate abundance at the oligotrophic stations in the the southern Yellow Sea. Heterotrophic flagellates did not increase while Synechococcus abundance.increased significantly. Therefore, there was no trophic cascade along the copepod-ciliate-flagellate food chain. Ciliates might be the dominant grazers of Synechococcus.
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