Analysis of the relationships between zooplankton and temperature-salinity based on RDA and GAMs model in coastal East China Sea
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摘要: 为进一步探讨东海近岸海域浮游动物季节演替与温盐的变化关系，根据2013年5月(春季)、8月(夏季)和12月(冬季)东海近岸海域3个航次的浮游动物调查资料，利用冗余分析模型分析了优势种与温盐的变化关系，广义加性模型分析了生物多样性参数与温盐之间的关系。结果表明：(1)优势种季节演替较为明显，由春季2种演替成夏季10种，再更替为冬季5种，春季和冬季优势种的种类和生态类群相对单一，夏季较为丰富。(2)优势种时空分布是综合多层环境因子的结果，如春季中华哲水蚤(Calanus sinicus)与表层盐度和底层盐度正相关，夏季与表层温度和底层温度负相关，冬季与表层盐度正相关；春季五角水母(Muggiaea atlantica)与底层温度、底层盐度呈现较好的正相关；夏季盐度是决定暖水性和高温高盐性优势种分布的关键因子，且呈现较强的正相关；冬季真刺唇角水蚤(Labidocera euchaeta)、强壮滨箭虫(Aidanosagitta crassa)与表层温度关系密切，并呈现较强的负相关。(3)春季和冬季香农–威纳指数多样性指数与纬度呈负相关。(4)温度和盐度对生物多样性参数影响显著，但季节上又呈现出差异。Abstract: In order to further explore the changes in relationships between seasonal succession of zooplankton and temperature-salinity in coastal East China sea, zooplankton survey data of three voyages in May (spring), August (summer) and December (winter) of 2013 were analyzed. Redundancy analysis model (RDA) was used to analyze the relationship between the changes of the dominant species and temperature-salinity, then the general algebraic modeling system (GAMs) was applied to analyze the relationship between temperature-salinity and parameters of the biological diversity. Results show that the dominant species’ succession changes seasonally from spring (2 kinds) to summer (10 kinds), then becomes 5 kinds in winter. Dominant species and the ecological groups are relatively single in spring and winter, but rich in summer. The temporal and spatial distribution of dominant species are the results of a comprehensive multi-level of environmental factors. For example, Calanus sinicus positively relates to surface salinity and bottom salinity in spring, negatively relates to surface temperature and bottom temperature in summer, positively relates to surface salinity in winter. Muggiaea atlantica presents better positive correlations with bottom temperature and bottom salinity in spring. What’s more, salinity is the key factor to determine the distribution of dominant species like warm water species and high temperature-salinity groups in summer, it also presents strong positive correlations. Labidocera euchaeta and Aidanosagitta crassa have strong negative correlations with surface temperature in winter. Shannon-Wiener diversity index negatively correlates with latitude in spring and winter. Biodiversity parameters are significantly affected by the temperature and salinity, but the influence presents differences seasonally.
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Key words:
- East China Sea /
- zooplankton /
- dominant species /
- biodiversity parameters /
- temperature /
- salinity
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图 1 采样站位
A. 东海北部近岸水域；B. 长江口及其邻近水域；C. 东海中南部近 岸水域
Fig. 1 Map of the sampling sites
A. Coastal waters of northern East China Sea; B. Changjiang River Estuary and adjacent waters; C. coastal waters of central and southern East China Sea
图 2 优势种与温盐的冗余分析模型（RDA）排序
Fig. 2 Redundancy analysis (RDA) ordination diagram of dominant species and temperature-salinity
图 4 平均温度、平均盐度与中华哲水蚤丰度分布关系
Fig. 4 The distribution relationship of C. sinicus abundance with average temperature and average salinity
图 5 香农–威纳多样性指数（H′）与纬度的线性关系
Fig. 5 The linear relationship of Shannon-Wiener diversity index（H′）and latitude
图 6 春季基于GAMs模型生物多样性参数与温盐的关系
Fig. 6 Relations in biodiversity parameters and temperature-salinity based on GAMs model in spring
图 7 夏季和冬季基于GAMs模型生物多样性参数与温盐的关系
Fig. 7 Relations in biodiversity parameters and temperature-salinity based on GAMs model in summer and winter
表 1 东海近岸海域浮游动物优势种的优势度（Y≥0.02）、平均丰度
Tab. 1 The dominance of dominant species （Y≥0.02）, average abundance in coastal waters of East China Sea
优势种 春季 夏季 冬季 种类 生态类群 Y 丰度均值/ind.·m–3 Y 丰度均值/ind.·m–3 Y 丰度均值/ind.·m–3 中华假磷虾 近岸低盐种 / / 0.03 5.72 0.03 0.53 强壮滨箭虫 近岸低盐种 / / / / 0.02 0.45 肥胖软箭虫 暖水种 / / 0.03 7.53 / / 百陶带箭虫 亚热带近海种 / / 0.05 11.20 0.03 0.58 纳噶带箭虫 亚热带近海种 / / 0.03 9.72 / / 中华哲水蚤 广温种 0.82 500.18 0.10 26.86 0.65 7.03 五角水母 暖水低盐种 0.04 39.70 / / / / 亚强次真哲水蚤 高温高盐种 / / 0.03 9.10 / / 精致真刺水蚤 暖水广盐种 / / 0.03 8.28 / / 平滑真刺水蚤 暖水种 / / 0.02 9.86 / / 真刺水蚤属 / / / 0.04 17.50 / / 真刺唇角水蚤 低盐种 / / 0.04 14.90 0.05 1.11 注：“/”表示非优势种。 
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表 2 东海近岸海域浮游动物优势种平均丰度区域分布（单位：ind./m3）
Tab. 2 Regional distribution of dominant species’ average abundance of zooplankton in coastal waters of East China Sea（unit: ind./m3）
春季 夏季 冬季 A B C A B C A B C 中华假磷虾 / / / 6.25 6.48 7.54 0.25 0.62 1.71 强壮滨箭虫 / / / / / / 0.52 1.35 0 肥胖软箭虫 / / / 3.10 13.50 8.23 / / / 百陶带箭虫 / / / 2.36 22.28 12.42 0.71 1.21 0.75 纳噶带箭虫 / / / 3.13 21.98 15.45 / / / 中华哲水蚤 662.63 184.75 645.29 3.45 41.45 57.47 6.14 10.12 3.43 五角水母 125.84 34.73 82.26 / / / / / / 亚强次真哲水蚤 / / / 0 6.93 22.80 / / / 精致真刺水蚤 / / / 2.40 8.08 17.07 / / / 平滑真刺水蚤 / / / 3.60 16.60 28.81 / / / 真刺水蚤属 / / / 0 26.25 57.29 / / / 真刺唇角水蚤 / / / 51.17 15.92 1.31 2.76 1.40 0 注：A. 东海北部近岸水域；B. 长江口及其邻近水域；C. 东海中南部近岸水域； “/”表示非优势种。
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表 3 RDA前两轴排序分析及蒙特卡洛检验结果
Tab. 3 Results by RDA ordination with the first two axis and Monte Carlo permutation test in herb layer
春季 夏季 冬季 RDA1 RDA 2 RDA 1 RDA 2 RDA 1 RDA 2 ST 0.512 0.727** –0.623* 0.219 –0.823** –0.522 SS 0.368 –0.769** 0.460 0.759** 0.070 –0.630* BT 0.709** 0.559 –0.702** 0.531 0.093 –0.727** BS 0.861** –0.374 0.772** 0.395 –0.179 –0.078 总被分析环境因子解释比例 43.21% 4.08% 15.10% 1.40% 14.77% 8.54% 注：ST表示表层温度；SS 表示表层盐度；BT 表示底层温度；BS 表示底层盐度；*表示 P<0.05；**表示 P<0.01。
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表 5 不同季节生物多样性参数与温盐关系GAMs模型分析结果
Tab. 5 Results of relations in biodiversity parameters and temperature-salinity based on GAMs model in different season
种类数S 多样性指数H′ 单纯度指数C 均匀度指数J′ 丰富度指数d 春季 环境因子 ST+SS+BT+BS ST+ BT+BS ST+ BT+BS ST+ SS+BS / AIC值 31.00 4.95 83.29 90.55 / 偏差解释比例/% 82.4 62.7 57.2 66.8 / 夏季 环境因子 / BS BS BS SS AIC值 / 34.19 50.31 61.65 29.35 偏差解释比例/% / 49.4 49.0 48.8 16.5 冬季 环境因子 ST+SS ST ST / ST AIC值 12.04 11.57 50.67 / 34.36 偏差解释比例/% 68.3 31.6 27.4 / 37.5 注：ST表示表层温度；SS表示表层盐度；BT表示底层温度；BS表示底层盐度。AIC赤池信息准则。“/”表示无数据。
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表 4 不同区域内的环境参数
Tab. 4 Environmental parameters in different areas
季节 区域 平均表温/℃ 平均表盐 平均底温/℃ 平均底盐 春季 A 13.7±0.78 30.2±1.07 12.6±1.27 30.6±0.95 B 15.7±0.88 26.4±3.06 15.5±1.72 30.7±3.56 C 17.0±0.74 29.3±1.45 17.9±1.39 31.3±2.38 夏季 A 26.3±0.75 29.9±0.83 26.5±0.76 29.8±0.67 B 25.7±0.65 28.3±5.00 23.3±2.28 31.0±4.58 C 26.2±0.70 32.0±1.21 25.2±2.47 32.6±1.21 冬季 A 8.3±0.52 30.1±0.88 8.6±0.52 30.2±0.68 B 11.8±2.59 29.8±2.95 17.0±2.78 29.4±4.15 C 12.9±2.27 29.5±1.72 13.3±2.38 29.7±1.82 注：A. 东海北部近岸水域; B. 长江口及邻近水域; C. 东海中南部近岸海域。
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