獐子岛海域浮游动物群落时空变化特征及其与环境因子之间的关系
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摘要
獐子岛海域地处北黄海长山群岛南端，距离辽东半岛南侧最近距离50公里，距离鸭绿江口150公里，是我国北方虾夷扇贝的重要的养殖区。该海域物理环境季节变化明显，夏季受黄海冷水团和鸭绿江冲淡水影响，冬季沿岸流和黄海暖流作用较大，是开展生物-物理耦合研究的理想海域
本论文利用2009年7月-2010年6月（除2010年1月与5月受恶劣天气影响未能登岛外）在獐子岛海域的周年逐月调查，分析了獐子岛海域浮游动物种类组成和丰度的周年变化情况，并分析了与环境因子的关系；通过与1959年普查时期浮游动物的周年变化数据对比，分析了该海域浮游动物群落的长期变化趋势。同时，重点分析了黄海冷水团对营养盐与叶绿素a浓度时空变化的影响，以及在中华哲水蚤种群结构和丰度周年变化过程中的作用主要研究结果如下：
在调查期间獐子岛海域大网出现浮游动物29种，浮游幼虫7种，共计36种，其中：中华哲水蚤和强壮箭虫为该海域全年优势种；细长脚（虫戎）在全年均有出现，主要为冬春季的优势种；双刺唇角水蚤、克氏纺锤水蚤、夜光虫、长腕类幼体同时为海域春季的优势种；鸟喙尖头溞为9月份优势种。该海域浮游动物年丰度和饵料生物量平均值分别为131.3ind/m3和217.5mg/m3，高峰值出现在6月，低值出现在8月。10月和11月观测到栉水母大量出现，并且10月份水母生物量与浮游动物丰度具有显著的负相关（P<0.05）。该海域生物多样性指数，丰富度指数最高值分别出现在4月（1.81）和10月（1.56）；均匀性指数较高，年平均值为0.49。与1959年普查数据相比，浮游动物优势种组成变动不大，丰度和多样性指数总体水平相近，但是2009年调查期间总丰度的波动幅度显著提高。
獐子岛海域夏秋季的营养盐和叶绿素浓度时空分布格局主要受黄海冷水团影响。2009年温跃层主要在7月到10的外海出现，表层盐度的降低出现在养殖海域的7月和8月。夏季（7月-9月）营养盐浓度在养殖区和外海都随深度的增加而不断增加，在11月和12月，各水层浓度分布较为平均。在水柱平均值上，除了铵盐，其它的营养盐都是从夏季到秋季不断增加。低于浮游植物最低吸收阈值的营养盐限制主要出现在夏季表层，但是化学计量学上的N限制在7月-12月一直存在。水柱叶绿素平均浓度7月-12月在外海要低于养殖区，8月10月养殖区叶绿素a浓度显著提高，但是在外海叶绿素a浓度变化较小。我们的结果显示营养盐限制主要出现在黄海冷水团存在的夏季表层，冲淡水的输入和黄海冷水团营养盐的积累能够促进浮游植物初级生产，虽然8月份降雨导致了浮游植物大量爆发，但是影响仅限于表层，虾夷扇贝的可利用食物并没有显著增加。与筏式养殖相比，贝类底播养殖并没有给生态环境造成很大影响，它能改变营养盐之间的比值，对叶绿素a浓度几乎没有影响。关于底播养殖对环境影响的调查有待于在大的区域进行更深入的调查研究。
黄海冷水团一直是优势种中华哲水蚤在中国沿海的度夏场所。在我们的周年调查中，中华哲水蚤的总丰度在10月份最低，在6月份最高；雌雄性比在2月份最高，8月份最低。水平分布的差异主要出现在黄海冷水团存在期间，在6月中华哲水蚤在养殖区的丰度为158.8ind/m3，在外海的丰度为532.1ind/m3；从7月到8月，黄海冷水团外中华哲水蚤的丰度从50.4ind/m3降低到1.9ind/m3，而在黄海冷水团内部中华哲水蚤丰度相似，分别为322.0ind/m3和324.4ind/m3。当9月份，黄海冷水团减弱的时候，中华哲水蚤分布较为平均。我们的结果显示黄海冷水团在春季延迟了中华哲水蚤的种群补充，但是在夏季保存了相当数量的度夏种群。成功度夏的种群在秋季扩散到近岸水域。除了低温，稳定的垂直结构也是中华哲水蚤度夏的关键条件。与黄海冷水团中部相比，边缘区也保存了相当丰度的中华哲水蚤度夏种群，但是种群结构（C5期个体所占比例及雌雄性比）明显不同。
The Zhangzi Island area, an important farming area for Patinopecten yessoensisin the Northern Yellow Sea, locates in the south part of the Changshan Archipelago,abouth50km from the south coast of Liaodong Penisulla and150km from the YaluRiver Estuary. Due to clear seasonal variation of physical environment, it is alsosuggested as an ideal place for biological-physical coupling studies.The physicalpattern is determined by the Yellow Sea Cold Water Mess (YSCWM) and Yalu Riverfreshwater discharge in summer and by coastal currents and the Yellow Sea WarmCurrent (YSWC) in winter.
In this study, monthly investgation was carried from July2009to June2010(except January and May for bad weather). Annual variation of zooplanktoncommunity was analyzed in relation to environmental conditions, includingtemperature regime, nutrient and Chlorophyll-a (Chl-a) concentrations. Long-termchange of zooplankton community was investigated through comparison withhistorical data in1959. Impacts of the YSCWM were emphasized on regimes ofnutrient and Chl a concentrations and population recruitment of Calanus sinicus,respectively.
During the investigation, a total of29zooplankton species and7planktoniclarvae were indentified. Calanus sinicus and Saggita crassa were observed asdominant species in all months. Themisto gracilipes was sampled in each month, butrecorded as dominant species only in winter and spring. In spring, Labidocerabipinnata, Acartia clausi, Noctiluca scintillans and Ophiopluteus larvae were alsorecorded as dominant species. Penetia avirostris was recorded as dominant speciesonly in September. Zooplankton abundance (non-jellyfish) and biomass were highestin June and lowest in August, with annual averages of131.3ind/m3and217.5mg/m3. A ctenophores Beroe bloom was observed in October and November. Beroe biomasscorrelated negatively and significantly with other zooplankton abundance in October(P<0.05). Zooplankton species diversity was highest in October (1.56), and richnessindex was highest in April (1.81). The average value of eveness index was0.49. Thedomiant species of zooplankton community and annual variation of total abundance inthis study was similar to those recorded in the former study in1959, indicating lowpossibility of regime shift. The difference in species composition may result frominter-annual variation in freshwater discharge and warm currents intrusion
Regimes of nutrient and Chl a concentration in summer and autumn wasdetermined mainly by the YSCWM. In2009, significant thermal stratificationpresented from July to October in open waters, and salinity decrease appeared in Julyand August in surface layers in mariculture area. Nutrient concentrations increasedwith depth in both areas in summer, but were similar through water column inNovember and December. On average, nutrient increased from summer to autumn inall components except ammonia. Nutrient concentrations lower than the minimumthresholds for phytoplankton growth presented only in upper layers in summer, butstoichiometric nitrogen limitation existed from July to December. Column averagedChl-a concentration was lower in open waters than in mariculture from July toDecember. It increased significantly in August and October in mariculture area, andwas less variable in open waters. Our results showed that nutrients limitation tophytoplankton growth presented mainly in upper layer in association withstratification caused by YSCWM in summer. Freshwater input and upwelling ofnutrients accumulated in YSCWM can stimulate phytoplankton production inmariculture area. Although phytoplankton bloom was induced by rainfall, littlecontribution was expected to food availability, as it was limited in surface layer.
The Yellow Sea Cold Water Mass (YSCWM) was suggested an over-summeringsite of the dominant copepod species Calanus sinicus in coastal China Seas. In ourstudy, total abundance was lowest in October and highest in June, and sex ratio washighest in February and lowest in August. Evident spatial difference in abundance wasobserved during existence of the YSCWM. In June, total abundance averaged in 158.8ind/m3at well-stratified stations, and532.1ind/m3at the other stations. FromJuly to August, abundance decreased from50.4to1.9ind/m3outside the YSCWM, butsimilarly high abundance of322.0and324.4ind/m3was recorded inside. When theYSCWM disappeared in September, C. sinicus distributed evenly over the study area.Our results indicate that existence of YSCWM retarded population recruitment inspring but preserved a cohort with considerable abundance in summer. Theover-summered population was transported to neritic waters in autumn. Besides lowtemperature, stable vertical structure is also essential condition for preservation ofover-summer population. Comparing to the central YSCWM area, C. sinicus cansurvive summer condition with considerable abundance in marginal area, butpopulation structure was completely different in terms of C5proportion and sex ratio.
本论文利用2009年7月-2010年6月（除2010年1月与5月受恶劣天气影响未能登岛外）在獐子岛海域的周年逐月调查，分析了獐子岛海域浮游动物种类组成和丰度的周年变化情况，并分析了与环境因子的关系；通过与1959年普查时期浮游动物的周年变化数据对比，分析了该海域浮游动物群落的长期变化趋势。同时，重点分析了黄海冷水团对营养盐与叶绿素a浓度时空变化的影响，以及在中华哲水蚤种群结构和丰度周年变化过程中的作用主要研究结果如下：
在调查期间獐子岛海域大网出现浮游动物29种，浮游幼虫7种，共计36种，其中：中华哲水蚤和强壮箭虫为该海域全年优势种；细长脚（虫戎）在全年均有出现，主要为冬春季的优势种；双刺唇角水蚤、克氏纺锤水蚤、夜光虫、长腕类幼体同时为海域春季的优势种；鸟喙尖头溞为9月份优势种。该海域浮游动物年丰度和饵料生物量平均值分别为131.3ind/m3和217.5mg/m3，高峰值出现在6月，低值出现在8月。10月和11月观测到栉水母大量出现，并且10月份水母生物量与浮游动物丰度具有显著的负相关（P<0.05）。该海域生物多样性指数，丰富度指数最高值分别出现在4月（1.81）和10月（1.56）；均匀性指数较高，年平均值为0.49。与1959年普查数据相比，浮游动物优势种组成变动不大，丰度和多样性指数总体水平相近，但是2009年调查期间总丰度的波动幅度显著提高。
獐子岛海域夏秋季的营养盐和叶绿素浓度时空分布格局主要受黄海冷水团影响。2009年温跃层主要在7月到10的外海出现，表层盐度的降低出现在养殖海域的7月和8月。夏季（7月-9月）营养盐浓度在养殖区和外海都随深度的增加而不断增加，在11月和12月，各水层浓度分布较为平均。在水柱平均值上，除了铵盐，其它的营养盐都是从夏季到秋季不断增加。低于浮游植物最低吸收阈值的营养盐限制主要出现在夏季表层，但是化学计量学上的N限制在7月-12月一直存在。水柱叶绿素平均浓度7月-12月在外海要低于养殖区，8月10月养殖区叶绿素a浓度显著提高，但是在外海叶绿素a浓度变化较小。我们的结果显示营养盐限制主要出现在黄海冷水团存在的夏季表层，冲淡水的输入和黄海冷水团营养盐的积累能够促进浮游植物初级生产，虽然8月份降雨导致了浮游植物大量爆发，但是影响仅限于表层，虾夷扇贝的可利用食物并没有显著增加。与筏式养殖相比，贝类底播养殖并没有给生态环境造成很大影响，它能改变营养盐之间的比值，对叶绿素a浓度几乎没有影响。关于底播养殖对环境影响的调查有待于在大的区域进行更深入的调查研究。
黄海冷水团一直是优势种中华哲水蚤在中国沿海的度夏场所。在我们的周年调查中，中华哲水蚤的总丰度在10月份最低，在6月份最高；雌雄性比在2月份最高，8月份最低。水平分布的差异主要出现在黄海冷水团存在期间，在6月中华哲水蚤在养殖区的丰度为158.8ind/m3，在外海的丰度为532.1ind/m3；从7月到8月，黄海冷水团外中华哲水蚤的丰度从50.4ind/m3降低到1.9ind/m3，而在黄海冷水团内部中华哲水蚤丰度相似，分别为322.0ind/m3和324.4ind/m3。当9月份，黄海冷水团减弱的时候，中华哲水蚤分布较为平均。我们的结果显示黄海冷水团在春季延迟了中华哲水蚤的种群补充，但是在夏季保存了相当数量的度夏种群。成功度夏的种群在秋季扩散到近岸水域。除了低温，稳定的垂直结构也是中华哲水蚤度夏的关键条件。与黄海冷水团中部相比，边缘区也保存了相当丰度的中华哲水蚤度夏种群，但是种群结构（C5期个体所占比例及雌雄性比）明显不同。
The Zhangzi Island area, an important farming area for Patinopecten yessoensisin the Northern Yellow Sea, locates in the south part of the Changshan Archipelago,abouth50km from the south coast of Liaodong Penisulla and150km from the YaluRiver Estuary. Due to clear seasonal variation of physical environment, it is alsosuggested as an ideal place for biological-physical coupling studies.The physicalpattern is determined by the Yellow Sea Cold Water Mess (YSCWM) and Yalu Riverfreshwater discharge in summer and by coastal currents and the Yellow Sea WarmCurrent (YSWC) in winter.
In this study, monthly investgation was carried from July2009to June2010(except January and May for bad weather). Annual variation of zooplanktoncommunity was analyzed in relation to environmental conditions, includingtemperature regime, nutrient and Chlorophyll-a (Chl-a) concentrations. Long-termchange of zooplankton community was investigated through comparison withhistorical data in1959. Impacts of the YSCWM were emphasized on regimes ofnutrient and Chl a concentrations and population recruitment of Calanus sinicus,respectively.
During the investigation, a total of29zooplankton species and7planktoniclarvae were indentified. Calanus sinicus and Saggita crassa were observed asdominant species in all months. Themisto gracilipes was sampled in each month, butrecorded as dominant species only in winter and spring. In spring, Labidocerabipinnata, Acartia clausi, Noctiluca scintillans and Ophiopluteus larvae were alsorecorded as dominant species. Penetia avirostris was recorded as dominant speciesonly in September. Zooplankton abundance (non-jellyfish) and biomass were highestin June and lowest in August, with annual averages of131.3ind/m3and217.5mg/m3. A ctenophores Beroe bloom was observed in October and November. Beroe biomasscorrelated negatively and significantly with other zooplankton abundance in October(P<0.05). Zooplankton species diversity was highest in October (1.56), and richnessindex was highest in April (1.81). The average value of eveness index was0.49. Thedomiant species of zooplankton community and annual variation of total abundance inthis study was similar to those recorded in the former study in1959, indicating lowpossibility of regime shift. The difference in species composition may result frominter-annual variation in freshwater discharge and warm currents intrusion
Regimes of nutrient and Chl a concentration in summer and autumn wasdetermined mainly by the YSCWM. In2009, significant thermal stratificationpresented from July to October in open waters, and salinity decrease appeared in Julyand August in surface layers in mariculture area. Nutrient concentrations increasedwith depth in both areas in summer, but were similar through water column inNovember and December. On average, nutrient increased from summer to autumn inall components except ammonia. Nutrient concentrations lower than the minimumthresholds for phytoplankton growth presented only in upper layers in summer, butstoichiometric nitrogen limitation existed from July to December. Column averagedChl-a concentration was lower in open waters than in mariculture from July toDecember. It increased significantly in August and October in mariculture area, andwas less variable in open waters. Our results showed that nutrients limitation tophytoplankton growth presented mainly in upper layer in association withstratification caused by YSCWM in summer. Freshwater input and upwelling ofnutrients accumulated in YSCWM can stimulate phytoplankton production inmariculture area. Although phytoplankton bloom was induced by rainfall, littlecontribution was expected to food availability, as it was limited in surface layer.
The Yellow Sea Cold Water Mass (YSCWM) was suggested an over-summeringsite of the dominant copepod species Calanus sinicus in coastal China Seas. In ourstudy, total abundance was lowest in October and highest in June, and sex ratio washighest in February and lowest in August. Evident spatial difference in abundance wasobserved during existence of the YSCWM. In June, total abundance averaged in 158.8ind/m3at well-stratified stations, and532.1ind/m3at the other stations. FromJuly to August, abundance decreased from50.4to1.9ind/m3outside the YSCWM, butsimilarly high abundance of322.0and324.4ind/m3was recorded inside. When theYSCWM disappeared in September, C. sinicus distributed evenly over the study area.Our results indicate that existence of YSCWM retarded population recruitment inspring but preserved a cohort with considerable abundance in summer. Theover-summered population was transported to neritic waters in autumn. Besides lowtemperature, stable vertical structure is also essential condition for preservation ofover-summer population. Comparing to the central YSCWM area, C. sinicus cansurvive summer condition with considerable abundance in marginal area, butpopulation structure was completely different in terms of C5proportion and sex ratio.
引文
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