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Article(id=1246833308584272067, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1246833307606995898, articleNumber=null, orderNo=null, doi=10.3969/j.issn.0253-4193.2019.08.007, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1521043200000, receivedDateStr=2018-03-15, revisedDate=1543766400000, revisedDateStr=2018-12-03, acceptedDate=null, acceptedDateStr=null, onlineDate=1775197825809, onlineDateStr=2026-04-03, pubDate=1566662400000, pubDateStr=2019-08-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1775197825809, onlineIssueDateStr=2026-04-03, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1775197825809, creator=13701087609, updateTime=1775197825809, updator=13701087609, issue=Issue{id=1246833307606995898, tenantId=1146029695717560320, journalId=1149651085930835976, year='2019', volume='41', issue='8', pageStart='1', pageEnd='140', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1775197825576, creator=13701087609, updateTime=1775200503343, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1246844539051332282, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1246833307606995898, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1246844539051332283, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1246833307606995898, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=72, endPage=84, ext={EN=ArticleExt(id=1246833308919816390, articleId=1246833308584272067, tenantId=1146029695717560320, journalId=1149651085930835976, language=EN, title=Analysis of the relationships between zooplankton and temperature-salinity based on RDA and GAMs model in coastal East China Sea, columnId=1243954927383462170, journalTitle=Haiyang Xuebao, columnName=Marine Biology, runingTitle=null, highlight=null, articleAbstract=

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.

, correspAuthors=Yunlong Wang, authorNote=null, correspAuthorsNote=null, copyrightStatement=Haiyang Xuebao, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Jieqing Yang, Yunrong Shi, Weimin Quan, Yunlong Wang), CN=ArticleExt(id=1246833310501069032, articleId=1246833308584272067, tenantId=1146029695717560320, journalId=1149651085930835976, language=CN, title=基于RDA与GAMs模型的东海近岸海域浮游动物与温盐关系, columnId=1243954927517679901, journalTitle=海洋学报, columnName=海洋生物, runingTitle=null, highlight=null, articleAbstract=

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。

, correspAuthors=王云龙, authorNote=null, correspAuthorsNote=
*王云龙(1964—)，男，研究员，主要从事海洋渔业生态的研究。E-mail：
, copyrightStatement=版权所有©《海洋学报》编辑部 2023
杨杰青，史赟荣，全为民，等. 基于RDA与GAMs模型的东海近岸海域浮游动物与温盐关系[J]. 海洋学报，2019，41(8)：72–84，doi:10.3969/j.issn.0253−4193.2019.08.007Yang Jieqing，Shi Yunrong，Quan Weimin, et al. Analysis of the relationships between zooplankton and temperature-salinity based on RDA and GAMs model in coastal East China Sea[J]. Haiyang Xuebao，2019, 41(8)：72–84，doi:10.3969/j.issn.0253−4193.2019.08.007
, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=EeGlEBmU/CAxybjKtDgnJw==, magXml=CDc8lZ5joaet1Kl/xxG20Q==, pdfUrl=null, pdf=NKS/7nCNcddQqG3g1cGyeQ==, pdfFileSize=1516015, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=h9zUx0hLQuYbPACO5IEZeg==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=E71aK7vIaQAF1ctS2mbLQQ==, mapNumber=null, authorCompany=null, fund=null, authors=

杨杰青（1990—），男，安徽省阜阳市人，研究方向为渔业生态学与环境保护。E-mail：

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杨杰青（1990—），男，安徽省阜阳市人，研究方向为渔业生态学与环境保护。E-mail：

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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

, figureFileSmall=KsT4j+TRBM1Ao4Ne6PscfA==, figureFileBig=ow8ZYII01edK8aNqcFQTsA==, tableContent=null), ArticleFig(id=1254506143598973769, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=CN, label=图1, caption=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

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The dominance of dominant species （Y≥0.02）, average abundance in coastal waters of East China Sea

, figureFileSmall=null, figureFileBig=null, tableContent=
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
), ArticleFig(id=1254506148577612657, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=CN, label=表1, caption=

东海近岸海域浮游动物优势种的优势度（Y≥0.02）、平均丰度

, figureFileSmall=null, figureFileBig=null, tableContent=
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
), ArticleFig(id=1254506148762162034, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=EN, label=Table 2, caption=

Regional distribution of dominant species’ average abundance of zooplankton in coastal waters of East China Sea（unit: ind./m3）

, figureFileSmall=null, figureFileBig=null, tableContent=
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
), ArticleFig(id=1254506148934128501, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=CN, label=表2, caption=

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）

, figureFileSmall=null, figureFileBig=null, tableContent=
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
), ArticleFig(id=1254506149194175352, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=EN, label=Table 3, caption=

Results by RDA ordination with the first two axis and Monte Carlo permutation test in herb layer

, figureFileSmall=null, figureFileBig=null, tableContent=
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%
), ArticleFig(id=1254506149433250683, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=CN, label=表3, caption=

RDA前两轴排序分析及蒙特卡洛检验结果

, figureFileSmall=null, figureFileBig=null, tableContent=
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%
), ArticleFig(id=1254506151136138108, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=EN, label=Table 5, caption=

Results of relations in biodiversity parameters and temperature-salinity based on GAMs model in different season

, figureFileSmall=null, figureFileBig=null, tableContent=
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
), ArticleFig(id=1254506151442322302, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=CN, label=表5, caption=

不同季节生物多样性参数与温盐关系GAMs模型分析结果

, figureFileSmall=null, figureFileBig=null, tableContent=
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
), ArticleFig(id=1254506151920472962, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=EN, label=Table 4, caption=

Environmental parameters in different areas

, figureFileSmall=null, figureFileBig=null, tableContent=
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
), ArticleFig(id=1254506152084050819, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833308584272067, language=CN, label=表4, caption=

不同区域内的环境参数

, figureFileSmall=null, figureFileBig=null, tableContent=
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
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基于RDA与GAMs模型的东海近岸海域浮游动物与温盐关系
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杨杰青 1 , 史赟荣 1 , 全为民 1 , 王云龙 1, *
海洋学报 | 海洋生物 2019,41(8): 72-84
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海洋学报 | 海洋生物 2019, 41(8): 72-84
基于RDA与GAMs模型的东海近岸海域浮游动物与温盐关系
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杨杰青1 , 史赟荣1, 全为民1, 王云龙1, *
作者信息
  • 1 中国水产科学研究院 东海水产研究所 农业农村部东海渔业资源开发利用重点实验室，上海 200090

    • 杨杰青（1990—），男，安徽省阜阳市人，研究方向为渔业生态学与环境保护。E-mail：

通讯作者:

*王云龙(1964—)，男，研究员，主要从事海洋渔业生态的研究。E-mail：
Analysis of the relationships between zooplankton and temperature-salinity based on RDA and GAMs model in coastal East China Sea
Jieqing Yang1 , Yunrong Shi1, Weimin Quan1, Yunlong Wang1, *
Affiliations
  • 1 Key Laboratory of East China Sea Fishery Resources Exploitation of Ministry of Agriculture and Rural Affairs, East China Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, Shanghai 200090, China
出版时间: 2019-08-25 doi: 10.3969/j.issn.0253-4193.2019.08.007
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摘要
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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。

东海  /  浮游动物  /  优势种  /  生物多样性参数  /  温度  /  盐度

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.

East China Sea  /  zooplankton  /  dominant species  /  biodiversity parameters  /  temperature  /  salinity
杨杰青, 史赟荣, 全为民, 王云龙. 基于RDA与GAMs模型的东海近岸海域浮游动物与温盐关系. 海洋学报, 2019 , 41 (8) : 72 -84 . DOI: 10.3969/j.issn.0253-4193.2019.08.007
Jieqing Yang, Yunrong Shi, Weimin Quan, Yunlong Wang. Analysis of the relationships between zooplankton and temperature-salinity based on RDA and GAMs model in coastal East China Sea[J]. Haiyang Xuebao, 2019 , 41 (8) : 72 -84 . DOI: 10.3969/j.issn.0253-4193.2019.08.007
正文
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1 引言
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东海是太平洋西岸一个受多种性质水系（沿岸径流、黄海冷水团、台湾暖流和黑潮暖流等）影响且较为开阔性的海区[1-2]，其水文环境复杂多变，地理区域跨度也较大（横跨温带和亚热带）[3-4]，因此浮游动物生态类群四季演替较为明显。浮游动物作为海洋次级生产者，对海洋物质循环和能量流动具有承上启下的作用[5-9]，另外作为海流或水团的指示种及水体环境变化的敏感类群[1, 10-11]，对于探索水团随季节变化特征及海洋生态环境质量状况等具有重要意义。
有关东海范围内浮游动物的生态研究已较为多见，多数集中在对浮游动物种类组成[12-14]、数量分布[15-16]、生态类群变化特征[17-18]、区域分布特征及多样性变化特征[4, 13, 15, 19-20]等研究上，虽有部分研究探讨了浮游动物群落结构与环境因子[21-24]的变化关系，但也只是局限于部分区域内（海湾、河口、保护区等）且分析方法较为单一，而有关整个东海近岸海域浮游动物优势种及生物多样性参数与环境因子的变化关系方面的研究还仍少见。冗余分析模型（Redundancy Analysis, RDA）是将样点投射到两条排序轴构成的二维平面上，通过样点的散集形态、在象限的分布等来反映研究区的特点，统计学上是通过原始变量与典型变量之间的相关性，分析引起原始变量变异的原因。广义加性模型（Generalized Additive Models, GAMs）是通过对自变量用非参数函数形式来拟合估计因变量和自变量之间的关系，其更加关注探测数据间的复杂关系，因其灵活性强，可控制季节趋势、长期趋势以及时间相关的混杂因素影响，目前已应用到许多学科领域中。
为了进一步探讨浮游动物优势种、生物多样性参数与温盐的关系，本文依据2013年5月、8月和12月3个航次的调查资料，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，以期为东海近岸海域浮游动物长期变化规律和生物多样性保护提供科学依据。
2 材料与方法
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2.1 调查时间及站位
调查范围位于东海北部近岸水域：32°N以北（含32°N，A1～A23站位，其中春季调查站位A1～A23，夏季A11～A23，冬季A11～A23）；长江口及其邻近水域：30°～31.5°N（B24～B44，春、夏和冬季调查站位均为B24～B44）；东海中南部近岸水域：29.5°N以南（含29.5°N，C45～C67站位；其中春季调查站位C45～C63，夏季C45～C56、C58～C63，冬季C45～C57）（图1）。
2.2 样品采集与分析
浮游动物样品采集与分析方法按《海洋调查规范》（GB/T12763.6–2007）[25]进行。针对近岸海域采用了浅水I型浮游生物网（配制网具规格为：网口内径50 cm, 网长145 cm，网囊网目为0.505 mm），由底层至表层进行垂直拖网一次，采集后的样品立即用福尔马林固定至最终溶液浓度为5%，带回实验室在体视显微镜下分析鉴定（包括种类、个数等）。环境参数（温度、盐度）采用YSI plus型水质分析仪现场测定。
2.3 数据处理与分析
2.3.1 优势度
浮游动物的优势度（Y）计算公式为
$Y=\left(n_{i} / N\right) \times f_{i} \text{，}$
式中，ni 为第i种的个体数，N为样品总个数，fi 为出现频率，以Y≥0.02为优势种的划分标准[20, 26]。
2.3.2 生物多样性指数
香农–威纳（Shannon-Wiener）多样性指数
$H^{\prime}=-\sum\limits_{i=1}^{S} P_{i} \log _{2} P_{i}\text{.}$
皮洛均匀度指数（Pielou evenness index）
$J^{\prime}=H^{\prime} / \log _{2} S\text{.}$
马加利夫丰富度指数（Margalef ’s index）
$d=(S-1) / \log _{2} N\text{.}$
单纯度指数
$C=\operatorname{sum}\left(n_{i} / N\right)^{2}\text{.}$
式（2）至式（5）中，S为该站记录种类数；Pi 为样品中第i种物种的个体比例，如样品总个数为N第i种个数为 ni ，则Pi = ni /N。
2.3.3 处理方法
为探讨优势种与温盐之间的相关性，并分析其分布受何种环境因子影响，通过物种样本做DCA分析，梯度长度的第一轴数值小于3.0，因此采用RDA线性分析，RDA排序图可较清晰地解释出二者之间的关系，图中温盐一般用带有箭头的线段表示，其箭头长短代表其在主轴中的作用大小，物种与温盐之间夹角的余弦值表示对应的相关性，夹角越小表示相关性越高，如果箭头同向，表示正相关，反向表示负相关，夹角接近直角，表示相关性很小。并采用蒙特卡洛检验环境因子与排序轴之间的相关性。因GAMs模型事先不需要进行相关性假设，又可探讨变量间的非单调、非线性关系，因此采用GAMs模型分析物种多样性指数与温盐之间的关系，根据多样性指数与温盐之间的显著性，筛选有效的环境因子进入分析模型，对模型进行赤池信息准则（Akaike’s Information Criterion，AIC）检验，当AIC数值最小时说明拟合效果最好，显著水平定为P<0.05。优势种丰度分布图采用Surfer 8绘制，香农–威纳多样性指数与纬度的关系图利用Excel 2010绘制，并用SPSS19.0线性回归分析其相关性。
3 结果
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3.1 优势种
春季共鉴定优势种(Y≥0.02)2种，分别为中华哲水蚤（Calanus sinicus）和五角水母（Muggiaea atlantica）（表1）。夏季优势种有10种，分别为中华假磷虾（Pseudeuphausia sinica）、肥胖软箭虫（Flaccisagitta enflata）、百陶带箭虫（Zonosagitta bedoti）、纳噶带箭虫（Z. nagae）、中华哲水蚤、亚强次真哲水蚤（Subeucalanus subcrassus）、精致真刺水蚤（Enchaeta concinna）、平滑真刺水蚤（E. plana）、真刺水蚤属（Euchaeta sp.）和真刺唇角水蚤（Labidocera euchaeta）。冬季优势种有5种，分别为中华假磷虾、强壮滨箭虫（Aidanisagitta crassa）、百陶带箭虫、中华哲水蚤和真刺唇角水蚤。其中，中华哲水蚤是3个季度共有优势种，优势种季节演替较为明显。
优势种季节和区域分布各具特色，其中中华哲水蚤平均丰度春季远高于夏季和冬季，而区域分布上春季东海北部近岸和中南部近岸海域要远高于长江口及其邻近海域（表2）。中华假磷虾、百陶带箭虫和真刺唇角水蚤平均丰度夏季远高于冬季，而区域分布上，中华假磷虾分布相对均匀，百陶带箭虫高值区在长江口及其邻近海域，真刺唇角水蚤高值区在东海北部近岸海域。
3.2 优势种与温盐的关系
春季总被分析环境因子解释比例为47.29%，其中RDA1解释比例为43.21%，RDA2解释比例为4.08%（表3），表4中底层温度（Bottom Temperature，BT）、底层盐度（Bottom Salinity，BS）与RDA1相关性较高（极显著，P<0.01），表层温度（Surface Temperature，ST）、表层盐度（Surface Salinity，SS）与RDA2密切相关（极显著，P<0.01）。夏季总被分析环境因子解释比例为16.50%，其中RDA1解释比例为15.10%，RDA2解释比例为1.40%，ST、BT、BS与RDA1排序轴相关性较高，其中与ST相关性达显著水平（P<0.05），与BT、BS达极显著水平（P<0.01），SS与RDA2排序轴具有较高的相关性（极显著，P<0.01）。冬季总被分析环境因子解释比例为24.31%，其中RDA1解释比例为14.77%，RDA2解释比例为8.54%，ST、BT与 RDA1排序轴相关性较高（极显著，P<0.01），SS与RDA2排序轴具有较高的相关性（P<0.05）（表3）。
如图2所示，温盐与物种之间的夹角余弦值越接近于1或–1，则表明相关性较密切，夹角余弦值越接近于0则表明相关性较小。春季五角水母与BT、BS密切相关，而中华哲水蚤与BS具有较为密切的关系。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；亚强次真哲水蚤与ST、SS和BS均具有较高的密切性，且与ST呈负相关，而与SS、BS呈正相关；真刺唇角水蚤与BT、BS密切相关，其中与BT正相关，与BS负相关；精致真刺水蚤、平滑真刺水蚤和真刺水蚤属对生态的适应性较为相似，与SS、BS正相关；中华哲水蚤、百陶带箭虫、纳噶带箭虫和肥胖软箭虫对温盐的适应性较为相似，均与ST、BT、BS相关，其中与ST正相关，与SS、BS负相关。冬季中华假磷虾与ST、BT关系较为密切，且呈正相关，中华哲水蚤、百陶带箭虫与SS关系较为密切，且呈正相关，强壮滨箭虫、真刺唇角水蚤与ST、BT负相关。
通过优势种与温盐的RDA排序关系，选择具有代表性的4个优势种（五角水母、中华哲水蚤、真刺唇角水蚤和强壮滨箭虫）探究了其区域分布特征。春季优势种五角水母主要分布于长江口以南的近岸海域，而冬季优势种真刺唇角水蚤和强壮滨箭虫主要分布于长江口以北（图3）。中华哲水蚤对温度的适应性较广（其温度的高丰度分布介于10～20℃之间），偏好稍高盐度的生境（盐度的高丰度分布介于27～32之间）（图4）。
3.3 香农–威纳多样性指数与纬度关系
春季、冬季香农–威纳多样性指数与纬度呈显著负相关，且达极显著水平（P<0.01），但夏季香农–威纳多样性指数与纬度相关性并不显著（P>0.05）（图5）。
3.4 多样性指数与温盐的关系
由表5可看出，春季决定浮游动物生物多样性指数是综合多层环境因子的结果，其中种类数S、多样性指数H′和单纯度指数C与ST、BT和BS相关，均匀度指数J′与ST、SS和BS相关，而夏季和冬季决定浮游动物生物多样性指数的环境因子相对较为单一，夏季与多样性指数H′、单纯度指数C 和均匀度指数J′相关的环境因子为BS，与丰富度指数d相关的环境因子为SS，冬季与种类数S、多样性指数H′、单纯度指数C 和丰富度指数d相关的环境因子主要是ST。
春季浮游动物种类数S与ST、SS、BT和BS之间的相关性各具差异，与ST先负相关后正相关（温度介于14～16℃时S较低），与BT相关性较为复杂，呈三峰变化，与SS负相关（SS越高，S越少），与BS相关性呈双峰变化（图6）。多样性指数H′与ST和BS的关系较为复杂，其相关性呈波动变化，与BT正相关。对比发现单纯度指数C和多样性指数H′分别与ST、BT和BS的相关性变化趋势刚好相反。
夏季BS小于20时，多样性指数H′和均匀度指数J′与BS正相关，20～28范围内多样性指数H′和均匀度指数J′与BS负相关，30～35范围内多样性指数H′和均匀度指数J′与BS又正相关，而单纯度指数C与BS的变化趋势恰恰相反。冬季种类数S、多样性指数H′和丰富度指数d与ST正相关，单纯度指数C与ST负相关。
4 讨论
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4.1 优势种分布与温盐的关系
东海近岸海域优势种季节演替较为明显，由春季2种演替成夏季10种，再更替为冬季的5种。春季和冬季优势种的种类和生态类群相对单一（表1），而夏季较为丰富。已有研究表明[21, 27-28]，浮游动物群落分布是综合各种环境因子影响的结果，其中温度、盐度是影响其分布的重要环境因子。由于不同种类的浮游动物对温盐耐受性的差异，因此其种类、数量和生态群落在区域及季节分布上又各显差异。
本研究发现，不同季节（春、夏、冬）的温度、盐度与中华哲水蚤丰度分布关系各有差异。中华哲水蚤对温度的适应性较广（其温度的高丰度分布介于10～20℃之间），偏好稍高盐度的生境（盐度的高丰度分布介于27～32之间）（图4）。林元烧和李松[29]在研究厦门港中华哲水蚤生活周期时发现，当水温高于24℃时中华哲水蚤便开始消失；Uye[30]研究认为实验室中中华哲水蚤胚胎发育水温上限为23℃；徐兆礼和李云[31]研究认为，中华哲水蚤生长最适温度为17.48℃。这种高温生境对中华哲水蚤丰度的限制在本研究中也得到证实，如夏季因调查海域平均水温处于一年中的最高值（ST>25℃，BT>21℃），大于其最适生长温度，中华哲水蚤分布与温度呈现较强的负相关；而冬、春两季水温相对较为适宜，温度又与其分布呈正相关（图2）。黄加祺和郑重[32]研究认为，中华哲水蚤最适盐度为30，本研究中华哲水蚤高丰度分布介于盐度27～32之间，因此也证实了中华哲水蚤偏好稍高盐度的生境。另外，生活史也决定着中华哲水蚤的丰度和区域分布，暖温种的中华哲水蚤主要分布于黄海和东海近海海域，春季是其大量繁殖的季节，因此在季节上具有明显的体现，如春季平均丰度最高（表1）。另外受长江口冲淡水影响（长江口及其邻近水域表层盐度较低），区域分布上也具有明显差异，如春季东海北部海域和中南部海域要远高于长江口及其邻近水域（表2）。
春、冬两季低盐性优势种（如五角水母、真刺唇角水蚤和强壮滨箭虫等）的分布主要取决于温度，如五角水母与BT呈现较好的正相关，真刺唇角水蚤、强壮滨箭虫与ST呈现较强的负相关（图2）。水文、气候等经过一年四季的轮回，春季整体水温较冬季有一定的提升，东海近岸海域水温呈现南高北低的特征（表4），长江口以南海域表层盐度整体不高，因此偏好暖水低盐性的五角水母[33]，其分布主要位于长江口以南的近岸海域，而偏好低温性的真刺唇角水蚤和强壮滨箭虫主要分布于长江口以北。这与丁峰元等[17]研究东海近海海域春季优势种（五角水母、强壮滨箭虫）群落分布较为一致。
夏季的盐度是决定暖水性和高温高盐性优势种（肥胖软箭虫、亚强次真哲水蚤、精致真刺水蚤和平滑真刺水蚤等）分布的关键因子，且盐度与其分布呈现较强的正相关。夏季外海黑潮暖流和南部台湾暖流给东海近岸带来了许多外海种和暖水种[34-36]，作为暖流所带来的优势种肥胖软箭虫、亚强次真哲水蚤、精致真刺水蚤和平滑真刺水蚤等，主要偏好高温高盐的环境。但因夏季整个东海近岸温度较高且相对均匀（表4），较适宜其生长繁殖，因此温度与其分布并未表现出显著的相关性。夏季沿岸径流（河口、海湾等）达一年中的最大值，近岸水体盐度分布不均匀（夏季盐度呈梯度变化，河口、海湾等近岸盐度低，远岸盐度高），因此这种环境格局也是盐度与其分布呈现较强正相关的最好解释。
4.2 香农–威纳多样性指数随纬度变化
香农–威纳多样性指数是基于物种数量反映群落种类多样性，种类越多、各物种个体均匀程度越高，即H′值愈大，群落所含的信息量愈大[37]。本研究发现，春、冬两季香农–威纳多样性指数与纬度呈负相关，但夏季相关性不显著。从种类数量方面可对春、冬两季香农–威纳多样性指数与纬度的负相关关系做出解释（低纬度浮游动物种类数多，高纬度种类数少）。
夏季香农–威纳多样性指数与纬度相关性不显著，这主要由香农–威纳多样性指数在纬度30.0°～31.0°N上偏高所引起（图5）。首先长江口与舟山渔场介于此纬度之间（图1），夏季该区域因台湾暖流和长江径流的汇合为该区域输送来多种生态类型的物种（如河口半咸水种、近岸低盐种、近岸暖水种、高温高盐种等）[21]，因此种类上相对较为丰富。其次长江口及邻近海域内平均均匀度指数（0.84）较其他两个区域的0.45与0.81都高，因此从数量均匀程度上也可以给予较好的解释。
4.3 生物多样性参数与温盐的关系
已有研究表明，温度和盐度可调控海域中浮游动物的种类组成、数量分布、群落结构及生物多样性等[10, 38]。而本研究也发现，温度和盐度对生物多样性参数也具有不同程度的影响。不同季节的温度和盐度与生物多样性参数之间的相关性也不相同，春季生物多样性参数是综合多层环境因子的结果，而冬、夏两季决定浮游动物生物多样性指数的环境因子相对较为单一。
春季整体水温还不高（表4），东海近岸受多种水团影响，其中东北部近岸仍受黄海冷水团控制[17]，水温较低，长江口及邻近水域属咸淡水交汇区，而东海中南部近岸海域也受微弱台湾暖流影响[36]，因此春季上层水域浮游动物分布呈斑块出现，分布不均匀，所以在种类数S、多样性指数H′上也具有一定体现（与ST的相关性波动变化）。研究表明浮游动物具有一定的垂直分布现象[39]，春季浮游动物多喜欢生活于下层水温高、盐度高的水域（图6），BT和BS越高其种类数S和多样性指数H′就越大，单纯度指数C越小（说明不单一）。夏季整体气温升高，表层水温达一年中最大值，黑潮暖流和台湾暖流也越加强劲[34-36]，因此较高的水温仅适宜于少数高温高盐种的生存，而限制了其他类群的存在，表层温盐并未起决定性作用，仅BS与多样性指数H′、单纯度指数C和均匀度指数J′显著相关，夏季受沿岸冲淡水及降雨量的影响，在咸淡水交汇区底层盐度20左右时其多样性指数H′和均匀度指数J′达最高值（图7）。冬季因整体水温最低，整个海区南北表层水温差异明显，多样性指数H′与纬度又极显著负相关，因此ST与浮游动物种类数S、多样性指数H′和丰富度指数d显著正相关，而与单纯度指数C显著负相关。
5 结论
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（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，如春季中华哲水蚤与SS和BS正相关，夏季与ST和BT负相关，冬季与SS正相关；春季五角水母与BT、BS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、强壮滨箭虫与ST关系密切，且呈现较强的负相关。
（3）春季和冬季香农–威纳多样性指数与纬度呈负相关，而夏季两者的相关性不显著。
（4）温度和盐度对生物多样性参数影响显著，又呈现出季节上的差异性。
基金
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  • 国家公益性行业（农业）科研专项经费项目（201303047）。
文献
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2019年第41卷第8期
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doi: 10.3969/j.issn.0253-4193.2019.08.007
  • 接收时间：2018-03-15
  • 首发时间：2026-04-03
  • 出版时间：2019-08-25
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  • 收稿日期：2018-03-15
  • 修回日期：2018-12-03
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国家公益性行业（农业）科研专项经费项目（201303047）。
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    1 中国水产科学研究院 东海水产研究所 农业农村部东海渔业资源开发利用重点实验室，上海 200090

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*王云龙(1964—)，男，研究员，主要从事海洋渔业生态的研究。E-mail：
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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