Article(id=1224796773516984625, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1224796769733726966, articleNumber=null, orderNo=null, doi=10.12284/hyxb2022180, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1649865600000, receivedDateStr=2022-04-14, revisedDate=1654531200000, revisedDateStr=2022-06-07, acceptedDate=null, acceptedDateStr=null, onlineDate=1769943906429, onlineDateStr=2026-02-01, pubDate=1664553600000, pubDateStr=2022-10-01, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1769943906429, onlineIssueDateStr=2026-02-01, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1769943906429, creator=13701087609, updateTime=1769943906429, updator=13701087609, issue=Issue{id=1224796769733726966, tenantId=1146029695717560320, journalId=1149651085930835976, year='2022', volume='44', issue='10', pageStart='1', pageEnd='192', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1769943905528, creator=13701087609, updateTime=1769996085810, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1225015629757956162, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1224796769733726966, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1225015629757956163, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1224796769733726966, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=100, endPage=108, ext={EN=ArticleExt(id=1224796773886083385, articleId=1224796773516984625, tenantId=1146029695717560320, journalId=1149651085930835976, language=EN, title=Simulation of spatio-temporal distribution of swordfish habitat in the western Indian Ocean based on maximum entropy model, columnId=1194652705852465724, journalTitle=Haiyang Xuebao, columnName=Article, runingTitle=null, highlight=null, articleAbstract=

Swordfish (Xiphias gladius) is a highly migratory fish whose habitat suitability is significantly influenced by the marine environment, and the prediction of its habitat using changes in the marine environment is of great scientific importance. In this study, we used the catch information of swordfish in the Chinese Indian Ocean Longline Fisheries Observer Data from 2017 to 2019 as species occurrence data, combined with the environmental data in the western Indian Ocean waters, including sea surface temperature (SST), sea surface height (SSH), chlorophyll a (Chl a) concentration, mixed layer depth (MLD), and sea surface salinity (SSS), the habitat suitability distribution of swordfish in the western Indian Ocean is simulated by using a maximum entropy model (MaxEnt). Model results show that: (1) the model has very high accuracy in simulating the habitat suitability distribution of swordfish in the western Indian Ocean, with AUC values greater than 0.9 in all seasons, and can be used to simulate the potential habitat suitability distribution of swordfish; (2) changes in the distribution of suitable habitat for swordfish in the study area are generally consistent with changes in the actual operational location, and the distribution of areas with high habitat suitability for swordfish is more concentrated in both the dry and rainy seasons, but the distribution range is greater in the wet season than in the dry season; (3) SST, SSS and MLD are important environmental factors affecting the habitat suitability distribution of swordfish in the western Indian Ocean. The optimum ranges of SST, SSS and MLD in the dry and rainy seasons are 25.8−31.6°C, 34.4−35.9 and 0.1−24.9 m, and 25.6−30.5°C, 34.8−36.4 and 13.1−54.1 m, respectively. The results of the study provide essential reference information for the sustainable use and scientific management of swordfish populations in the western Indian Ocean.

, correspAuthors=Xuefang Wang, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright © 2022 Pratacultural Science. All rights reserved., 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=Wei Tang, Xuefang Wang, Feng Wu, Yuan Li), CN=ArticleExt(id=1224796775387644267, articleId=1224796773516984625, tenantId=1146029695717560320, journalId=1149651085930835976, language=CN, title=基于最大熵模型模拟西印度洋剑鱼栖息地的时空分布, columnId=1149698756456657529, journalTitle=海洋学报, columnName=论文, runingTitle=null, highlight=null, articleAbstract=

剑鱼(Xiphias gladius)是一种高度洄游性鱼类,其迁徙和栖息地利用受海洋环境影响明显,理解其空间分布格局形成的机制对于资源的养护和管理具有重要意义。本研究利用2017−2019年中国印度洋延绳钓渔业观察员数据中剑鱼的渔获物信息作为物种出现数据,结合西印度洋海域的海表温度、海面高度、叶绿素a浓度、混合层深度、海表盐度等环境数据,采用最大熵模型对剑鱼的栖息地适宜性分布进行了模拟。结果表明:(1)模型对印度洋西部剑鱼栖息地适宜性分布的模拟精度非常高,各个季节受试者工作特征曲线的曲线下面积都大于0.9,可用于模拟剑鱼潜在的栖息地适宜性分布;(2)研究区域内剑鱼适宜栖息地分布变化与实际作业位置变动基本一致,干季和湿季剑鱼栖息地高适宜性的区域分布都较为集中,但湿季分布范围要大于干季;(3)海表温度、海表盐度和混合层深度是影响西印度洋剑鱼栖息地适宜性分布的重要环境因子,在干季和湿季的最适范围分别为25.8~31.6℃、34.4~35.9、0.1~24.9 m和25.6~30.5℃、34.8~36.4、13.1~54.1 m。研究结果可为西印度洋海域剑鱼种群的可持续利用和科学管理提供必要参考信息。

, correspAuthors=王学昉, authorNote=null, correspAuthorsNote=
王学昉(1983-),男,副教授,硕导,研究方向为海洋渔业资源监测与栖息地评估。E-mail:
, copyrightStatement=版权所有©《海洋学报》编辑部 2022, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=47F/8873MAX3TdP3JdBn3A==, magXml=BRNdK1BqWyfl7ypT4X86Xw==, pdfUrl=null, pdf=T0WraLDwaMi3fJOPXpVstA==, pdfFileSize=2398952, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=MYVdP1c+3JPKsRS5rQCKgA==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=/IzSRwKuo8TLsG9Qc1ke0A==, mapNumber=null, authorCompany=null, fund=null, authors=

唐未(1998-),男,四川省巴中市人,研究方向为鱼类栖息地评估。E-mail:

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唐未(1998-),男,四川省巴中市人,研究方向为鱼类栖息地评估。E-mail:

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唐未(1998-),男,四川省巴中市人,研究方向为鱼类栖息地评估。E-mail:

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a. Dry season in 2017; b. dry season in 2018; c. dry season in 2019; d. rainy season in 2017; e. rainy season in 2018; f. rainy season in 2019

, figureFileSmall=75oQpsxBti+gHAP9COkoPg==, figureFileBig=EzryZ25hDbksGRVMjU0KMQ==, tableContent=null), ArticleFig(id=1225368254600884765, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=CN, label=图2, caption=2017−2019年西印度洋剑鱼实际出现点与潜在栖息地的分布

a. 2017年干季;b. 2018年干季;c. 2019年干季;d. 2017年湿季; e. 2018年湿季; f. 2019年湿季

, figureFileSmall=75oQpsxBti+gHAP9COkoPg==, figureFileBig=EzryZ25hDbksGRVMjU0KMQ==, tableContent=null), ArticleFig(id=1225368254705742370, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=EN, label=Fig. 3, caption=Distribution of swordfish habitat suitability index in the western Indian Ocean from 2017 to 2019

a. Dry season in 2017; b. dry season in 2018; c. dry season in 2019; d. rainy season in 2017; e. rainy season in 2018; f. rainy season in 2019

, figureFileSmall=wA8bulvNyQAnRonoSN0PuA==, figureFileBig=wSJ2rSzsnfpXVR84CCGReA==, tableContent=null), ArticleFig(id=1225368254798017063, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=CN, label=图3, caption=2017−2019年西印度洋剑鱼栖息地适宜性指数分布

a. 2017年干季;b. 2018年干季;c. 2019年干季;d. 2017年湿季; e. 2018年湿季; f. 2019年湿季

, figureFileSmall=wA8bulvNyQAnRonoSN0PuA==, figureFileBig=wSJ2rSzsnfpXVR84CCGReA==, tableContent=null), ArticleFig(id=1225368254869320234, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=EN, label=Fig. 4, caption=Comparison of habitat suitability index for swordfish in the western Indian Ocean during the dry and rainy seasons from 2017 to 2019, figureFileSmall=7yzl/YgZYHM7vczik95Apg==, figureFileBig=MsSM/z/xwJDn2OngN3w2Vg==, tableContent=null), ArticleFig(id=1225368254965789230, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=CN, label=图4, caption=2017−2019年干季和湿季西印度洋剑鱼栖息地适宜性指数比较, figureFileSmall=7yzl/YgZYHM7vczik95Apg==, figureFileBig=MsSM/z/xwJDn2OngN3w2Vg==, tableContent=null), ArticleFig(id=1225368255062258227, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=EN, label=Fig. 5, caption=Response curves of main environmental factors to swordfish habitat suitability index, figureFileSmall=phb5hB6BFQ9RFiwVBX99kg==, figureFileBig=Z/CeWwC52wy+C2d6D7Vayw==, tableContent=null), ArticleFig(id=1225368255125172789, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=CN, label=图5, caption=主要环境因子对剑鱼栖息地适宜性指数的响应曲线, figureFileSmall=phb5hB6BFQ9RFiwVBX99kg==, figureFileBig=Z/CeWwC52wy+C2d6D7Vayw==, tableContent=null), ArticleFig(id=1225368255221641785, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=EN, label=Table 1, caption=

Main parameters for model evaluation and test

, figureFileSmall=null, figureFileBig=null, tableContent=
2017年干季2017年湿季2018年干季2018年湿季2019年干季2019年湿季
训练数据(AUC值)0.965 70.93430.987 20.933 90.976 90.969 4
测试数据 (AUC值)0.965 40.92470.981 80.925 30.965 60.965 2
AUC值标准差0.005 40.01360.009 80.014 40.011 00.009 4
ESS值0.251 20.35090.349 70.323 10.285 10.154 0
遗漏率/%9.814.25.615.05.69.4
), ArticleFig(id=1225368255288750657, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=CN, label=表1, caption=

模型预测评价及主要验证参数

, figureFileSmall=null, figureFileBig=null, tableContent=
2017年干季2017年湿季2018年干季2018年湿季2019年干季2019年湿季
训练数据(AUC值)0.965 70.93430.987 20.933 90.976 90.969 4
测试数据 (AUC值)0.965 40.92470.981 80.925 30.965 60.965 2
AUC值标准差0.005 40.01360.009 80.014 40.011 00.009 4
ESS值0.251 20.35090.349 70.323 10.285 10.154 0
遗漏率/%9.814.25.615.05.69.4
), ArticleFig(id=1225368255372636742, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=EN, label=Table 2, caption=

The contribution rate (%) of environmental factors in the seasonal maximum entropy model from 2017 to 2019

, figureFileSmall=null, figureFileBig=null, tableContent=
环境因子2017年
干季
2017年
湿季
2018年
干季
2018年
湿季
2019年
干季
2019年
湿季
SST41.3925.8732.3446.7654.9346.85
SSS17.7533.6715.1831.6720.7313.69
SSH18.7717.846.403.553.183.23
Chl a浓度1.991.7128.449.0614.7828.07
MLD20.1020.9117.648.966.388.16
), ArticleFig(id=1225368255443939913, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=CN, label=表2, caption=

2017−2019年各季最大熵模型中环境因子的贡献率 (%)

, figureFileSmall=null, figureFileBig=null, tableContent=
环境因子2017年
干季
2017年
湿季
2018年
干季
2018年
湿季
2019年
干季
2019年
湿季
SST41.3925.8732.3446.7654.9346.85
SSS17.7533.6715.1831.6720.7313.69
SSH18.7717.846.403.553.183.23
Chl a浓度1.991.7128.449.0614.7828.07
MLD20.1020.9117.648.966.388.16
), ArticleFig(id=1225368255523631695, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=EN, label=Table 3, caption=

The seasonal result score of Jackknife test from 2017 to 2019

, figureFileSmall=null, figureFileBig=null, tableContent=
2017年
干季
2017年
湿季
2018年
干季
2018年
湿季
2019年
干季
2019年
湿季
不包含MLD2.2381.4712.2211.5342.4332.485
不包含Chl a浓度2.2591.5872.8521.4102.0642.394
不包含SSH2.2441.4782.6791.5372.3792.491
不包含SSS2.1151.2022.4360.9702.3122.054
不包含SST2.1601.5812.5641.2052.2472.285
只包含MLD1.4160.7841.4620.6790.9850.959
只包含Chl a浓度0.6900.2061.1890.4071.0630.161
只包含SSH1.0790.8770.3670.5660.6361.015
只包含SSS1.0440.8051.5120.5470.9980.874
只包含SST1.1450.4200.9240.5741.1281.338
), ArticleFig(id=1225368255599129169, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1224796773516984625, language=CN, label=表3, caption=

2017−2019年各季Jackknife检验结果得分

, figureFileSmall=null, figureFileBig=null, tableContent=
2017年
干季
2017年
湿季
2018年
干季
2018年
湿季
2019年
干季
2019年
湿季
不包含MLD2.2381.4712.2211.5342.4332.485
不包含Chl a浓度2.2591.5872.8521.4102.0642.394
不包含SSH2.2441.4782.6791.5372.3792.491
不包含SSS2.1151.2022.4360.9702.3122.054
不包含SST2.1601.5812.5641.2052.2472.285
只包含MLD1.4160.7841.4620.6790.9850.959
只包含Chl a浓度0.6900.2061.1890.4071.0630.161
只包含SSH1.0790.8770.3670.5660.6361.015
只包含SSS1.0440.8051.5120.5470.9980.874
只包含SST1.1450.4200.9240.5741.1281.338
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基于最大熵模型模拟西印度洋剑鱼栖息地的时空分布
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唐未 1 , 王学昉 1, 2, 3, 4, * , 吴峰 1, 2, 3, 4 , 李渊 5
海洋学报 | 论文 2022,44(10): 100-108
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海洋学报 | 论文 2022, 44(10): 100-108
基于最大熵模型模拟西印度洋剑鱼栖息地的时空分布
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唐未1 , 王学昉1, 2, 3, 4, * , 吴峰1, 2, 3, 4, 李渊5
作者信息
  • 1.上海海洋大学 海洋科学学院,上海 201306
  • 2.国家远洋渔业工程技术研究中心,上海 201306
  • 3.大洋渔业资源可持续开发教育部重点实验室,上海 201306
  • 4.农业农村部大洋渔业资源环境科学观测实验站,上海 201306
  • 5.自然资源部第三海洋研究所,福建 厦门 361005
  • 唐未(1998-),男,四川省巴中市人,研究方向为鱼类栖息地评估。E-mail:

通讯作者:

王学昉(1983-),男,副教授,硕导,研究方向为海洋渔业资源监测与栖息地评估。E-mail:
Simulation of spatio-temporal distribution of swordfish habitat in the western Indian Ocean based on maximum entropy model
Wei Tang1 , Xuefang Wang1, 2, 3, 4, * , Feng Wu1, 2, 3, 4, Yuan Li5
Affiliations
  • 1. College of Marine Sciences, Shanghai Ocean University, Shanghai 201306, China
  • 2. National Distant-water Fisheries Engineering Research Center, Shanghai 201306, China
  • 3. The Key Laboratory of Sustainable Exploitation of Oceanic Fisheries Resources, Ministry of Education, Shanghai 201306, China
  • 4. Scientific Observing and Experimental Station of Oceanic Fishery Resources, Ministry of Agriculture and Rural Affairs, Shanghai 201306, China
  • 5. Third Institute of Oceanography, Ministry of Natural Resources, Xiamen 361005, China
出版时间: 2022-10-01 doi: 10.12284/hyxb2022180
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剑鱼(Xiphias gladius)是一种高度洄游性鱼类,其迁徙和栖息地利用受海洋环境影响明显,理解其空间分布格局形成的机制对于资源的养护和管理具有重要意义。本研究利用2017−2019年中国印度洋延绳钓渔业观察员数据中剑鱼的渔获物信息作为物种出现数据,结合西印度洋海域的海表温度、海面高度、叶绿素a浓度、混合层深度、海表盐度等环境数据,采用最大熵模型对剑鱼的栖息地适宜性分布进行了模拟。结果表明:(1)模型对印度洋西部剑鱼栖息地适宜性分布的模拟精度非常高,各个季节受试者工作特征曲线的曲线下面积都大于0.9,可用于模拟剑鱼潜在的栖息地适宜性分布;(2)研究区域内剑鱼适宜栖息地分布变化与实际作业位置变动基本一致,干季和湿季剑鱼栖息地高适宜性的区域分布都较为集中,但湿季分布范围要大于干季;(3)海表温度、海表盐度和混合层深度是影响西印度洋剑鱼栖息地适宜性分布的重要环境因子,在干季和湿季的最适范围分别为25.8~31.6℃、34.4~35.9、0.1~24.9 m和25.6~30.5℃、34.8~36.4、13.1~54.1 m。研究结果可为西印度洋海域剑鱼种群的可持续利用和科学管理提供必要参考信息。

西印度洋  /  剑鱼  /  栖息地模拟  /  最大熵模型  /  仅出现数据

Swordfish (Xiphias gladius) is a highly migratory fish whose habitat suitability is significantly influenced by the marine environment, and the prediction of its habitat using changes in the marine environment is of great scientific importance. In this study, we used the catch information of swordfish in the Chinese Indian Ocean Longline Fisheries Observer Data from 2017 to 2019 as species occurrence data, combined with the environmental data in the western Indian Ocean waters, including sea surface temperature (SST), sea surface height (SSH), chlorophyll a (Chl a) concentration, mixed layer depth (MLD), and sea surface salinity (SSS), the habitat suitability distribution of swordfish in the western Indian Ocean is simulated by using a maximum entropy model (MaxEnt). Model results show that: (1) the model has very high accuracy in simulating the habitat suitability distribution of swordfish in the western Indian Ocean, with AUC values greater than 0.9 in all seasons, and can be used to simulate the potential habitat suitability distribution of swordfish; (2) changes in the distribution of suitable habitat for swordfish in the study area are generally consistent with changes in the actual operational location, and the distribution of areas with high habitat suitability for swordfish is more concentrated in both the dry and rainy seasons, but the distribution range is greater in the wet season than in the dry season; (3) SST, SSS and MLD are important environmental factors affecting the habitat suitability distribution of swordfish in the western Indian Ocean. The optimum ranges of SST, SSS and MLD in the dry and rainy seasons are 25.8−31.6°C, 34.4−35.9 and 0.1−24.9 m, and 25.6−30.5°C, 34.8−36.4 and 13.1−54.1 m, respectively. The results of the study provide essential reference information for the sustainable use and scientific management of swordfish populations in the western Indian Ocean.

the western Indian Ocean  /  Xiphias gladius  /  habitat simulation  /  maximum entropy model (MaxEnt)  /  presence-only data
唐未, 王学昉, 吴峰, 李渊. 基于最大熵模型模拟西印度洋剑鱼栖息地的时空分布. 海洋学报, 2022 , 44 (10) : 100 -108 . DOI: 10.12284/hyxb2022180
Wei Tang, Xuefang Wang, Feng Wu, Yuan Li. Simulation of spatio-temporal distribution of swordfish habitat in the western Indian Ocean based on maximum entropy model[J]. Haiyang Xuebao, 2022 , 44 (10) : 100 -108 . DOI: 10.12284/hyxb2022180
剑鱼(Xiphias gladius)是一种高度洄游性鱼类,广泛分布于三大洋热带和亚热带海域[1],常被金枪鱼延绳钓商业船队所兼捕[2]。除经济价值以外,剑鱼作为一种海洋顶端捕食者在中上层生态系统中发挥着重要作用[3],探究其迁移规律和栖息地利用的基础研究一直备受关注[4-6]
物种分布模型(Species Distribution Model, SDM)是渔业生态学家和管理者评估和管理海洋生物栖息地和分布的重要工具[7],已有数个研究使用栖息地适宜性指数(Habitat Suitability Index, HSI)模型[8]、广义加性模型(Generalized Additive Model, GAM) [4]、广义线性模型(General Linear Model, GLM)[9]和多物种矢量自回归时空模型(Multi-species Vector-Autoregressive Spatio-Temporal Model, VAST)[10]等模拟预测剑鱼在不同海域的栖息地适宜性。这些方法有的需要基于丰富的专家经验进行判断,有的要求高质量的物种出现数据作为支撑,而剑鱼作为兼捕渔获物,其生态学信息和丰度数据在印度洋海域都十分匮乏,导致需要使用更为合适的建模方法进行栖息地量化的尝试[11-12]
最大熵模型(Maximum Entropy Model, MaxEnt)是基于最大熵原理预测物种地理空间分布的模型[13],是一种基于“仅出现”预测“未知分布”的机器学习方法[14]。MaxEnt只需要使用“仅出现”数据(“presence-only” 数据),就能利用高效简明的数学方法来模拟物种的栖息地适宜性,同时具有一种明确的“正则化”机制来防止模型过拟合,在降低数据精度要求的前提下,仍具有很高的预测精度[14-15]。目前MaxEnt已被广泛应用于海洋鱼类、鲸豚类、底栖生物、珊瑚等物种分布的预测研究[16-19]
因此,本研究将利用MaxEnt对西印度洋剑鱼栖息地适宜性进行建模模拟,以获取该海域剑鱼栖息地适宜性的时空变化特征和海洋环境关键影响因素,旨在为西印度洋海域剑鱼种群的可持续利用和科学管理提供必要参考信息。
研究区域为西印度洋,范围为30°N~40°S,30°~80°E(图1)。在该区域内,营养物质丰富,浮游生物大量繁殖[20],是剑鱼重要的产卵场和栖息场所[21],但海洋环境受到季风和洋流的强烈影响,温度、盐度和混合层深度等带有显著的季节性变化[22-25]。其鲜明的环境特征造就明显的干湿季节,湿润季节通常从5月底开始,持续到11月;干旱季节从11月底一直持续到翌年4月[26]
本研究选择2017−2019年中国印度洋金枪鱼延绳钓渔业观察员数据中记录的1 017条剑鱼渔获作为物种出现数据,包括作业年份、月份、捕捞位置等。
其他海域的研究表明,剑鱼的时空分布主要受海表温度(Sea Surface Temperature, SST)、海面高度(Sea Surface Height, SSH)、叶绿素a(chlorophyll a, Chl a)浓度、混合层深度(Mixed Layer Depth, MLD)和海表盐度(Sea Surface Salinity, SSS)等海洋环境因素的影响[4-6, 8-10]。因此本研究拟采用这5种海洋环境因子用于构建栖息地模型。环境数据均来自于哥白尼海洋环境监测服务中心(Copernicus-Marine Environment Monitoring Service, CMEMS) (https://resources.marine.copernicus.eu)。其中混合层深度数据的原始数据时间分辨率为月,空间分辨率为(1/12)°;海表温度、海表盐度、叶绿素a浓度和海面高度原始数据时间分辨率为月,空间分辨率为(1/4)°。
输入到MaxEnt的各数据图层必须是同一分辨率以便匹配使用。因此使用ArcMap10.4软件将环境数据输入进行栅格叠加并求平均值,全部转换成(1/4)°×(1/4)°的空间分辨率,数据的空间范围为30°N~40°S,30°~80°E。时间上按照各年干湿季节为间隔转换。利用ArcMap10.4软件将环境数据图层转换为ASCII栅格数据。将剑鱼渔获位置的经纬度数据转换为 csv格式保存。
使用软件MAXENT3.4.1(http://biodiversityinformatics. amnh.org/open_source/maxent/),随机将剑鱼样本中80%的出现点设置为训练集,剩余20%的出现点作为测试集,重复计算10次,以消除随机性,并去除重复数[15, 17, 19],结果以Logistic格式输出,绘制响应曲线,使用刀切法分析环境因子与剑鱼栖息地适宜性的相互关系。以训练模型灵敏度与稳定性相等(Equal Training Sensitivity and Specificity, ESS)时的剑鱼栖息地适宜性值为阈值,衡量测试点的遗漏率[27],区分适宜栖息地和非适宜栖息地,从而检验剑鱼出现点未被模型预测到的比率。
使用受试者工作特征曲线(Receiver Operating Characteristic Curve, ROC)的曲线下面积(Area Under Curve, AUC)的大小作为评估MaxEnt预测准确度的指标[14]。在物种分布模型预测准确度评估时,一般考虑范围为0.5~1的AUC值,0.5为随机模型的AUC值,AUC值越大则表明模型预测的准确度越高,通常AUC值大于0.7时,该模型即可当作良好的预测模型[28]
通过模型中的Jackknife检验模块,分析不同时期各环境因子对剑鱼栖息地适宜性的贡献率,通过比较仅存在某环境因子和不存在该环境因子时的得分大小,来评估其对剑鱼栖息地适宜性的重要性。
根据MaxEnt输出的(1/4)°×(1/4)°栅格点上的栖息地适宜性数值,采用ArcMap10.4软件绘制模型模拟的西印度洋剑鱼栖息地适宜性分布地图。
表1所示,研究期内各季的AUC值均显著大于0.5,表明MaxEnt预测结果显著优于随机分布模型。同时每个季节的训练数据和测试数据得到的AUC值都高于0.9,说明采用的环境变量对剑鱼的栖息地选择具有重要作用,得到的栖息地适宜性模拟结果具有很好的区分能力,模型模拟的结果优异,可用于西印度洋剑鱼栖息地适宜性分布的研究。
ESS 值的结果表明,大部分剑鱼出现点都分布在模型识别的适宜区域内(图2),但湿季的模型遗漏率均值(12.9%)高于干季的模型遗漏率均值(7%,表1)。
图2展示了模型模拟的西印度洋海域剑鱼潜在栖息地的时空分布变化状况,可以发现湿季和干季的适宜区域范围存在着显著的差别:干季都集中于马达加斯加北面和东面(图2a图2c),而湿季则明显向北半球转移,到达索马里东部,扩散至阿拉伯海区域(图2d图2f),且总体分布范围要大于干季。剑鱼栖息地适宜性指数的空间分布也呈现出相似格局,2017−2018年湿季分布范围大于干季且向南北分散(图3d图3e),但2019年湿季在30ºS水域则未出现适宜性区域(图3f),而所有情景下高适宜性的区域(HSI>0.7)都表现出较为集中的特征(图3)。将剑鱼出现点所对应的栅格值提取并进行统计发现:研究期内湿季的HSI总体水平要高于干季,差距不大且各年间变化不一;但2018年干季是一个例外,其均值约为0.75,显著高于其他各季0.55左右的水平(图4)。
从环境因子在各季MaxEnt贡献率中可发现(表2),SST的贡献率最高,平均为41.36%,2019年干季贡献率最高,达到54.93%。SSS的贡献率位居其次,平均为22.12%,2017年湿季贡献率最高,达到33.67%。SSH的平均贡献率最低,仅为8.83%,但在不同年份和季节的差异很大。
根据Jackknife检验可知(表3),不包含SST和SSS的平均得分低于不包含Chl a浓度、SSH 和MLD 3项。在只包含某个环境因子的得分上,只包含SSS和MLD 的平均得分要高于只包含SST、Chl a浓度和SSH 的得分。因此,研究期内SST是影响西印度洋剑鱼栖息地适宜性变化的首要影响因子,SSS和MLD是次要影响因素。
本研究选取MaxEnt模拟的研究区域内所有栅格HSI最大的前10%作为剑鱼的最适宜栖息地,计算得到干季研究区域内剑鱼最适宜栖息地的SST为25.8~31.6℃,SSS为34.4~35.9,MLD为0.1~24.9 m;湿季研究区域内剑鱼最适宜栖息地的SST为25.6~30.5℃,SSS为34.8~36.4,MLD为13.1~54.1 m。从剑鱼栖息地适宜性变化的主要环境因子的影响曲线可知(图5),湿季与干季剑鱼栖息地SSS的最适范围差异不大,而湿季剑鱼对SST的耐受范围要更宽一些,且倾向于更深的MLD环境。
SDM作为在生态学、生物地理学和分类学等方面用来预测物种潜在分布的重要和常用工具[29-31],在许多领域都发挥着重要作用,随着技术发展也开发了许多不同方法,如生物气候分析系统模型(Bioclimate Analysis System Model, BIOCLIM)[32]、广义线性模型[33]、广义加性模型[34]、人工神经网络(Artificial Neural Networks, ANN)[35]和随机森林(Random Forests, RF)[36]等,这些模型都需要出现/非出现数据作为物种的发生记录[37]
但是对于大量的海洋生物类群而言,很难在复杂的海洋环境中获取可靠的非出现数据[38]。因此,像MaxEnt这类利用仅出现或出现/伪缺失信息的算法可能非常有用[14-15]。特别当数据量有限时,MaxEnt就是一种良好的替代方法[14],甚至在某些时候表现更好[39-40]:如Tittensor等[18]通过MaxEnt和生态位因子分析(Environmental Niche Factor Analysis, ENFA)两种方法对全球范围的石珊瑚栖息地适宜性进行模拟,结果表明MaxEnt的效果一直优于ENFA;Monk等[19]使用包括MaxEnt、ENFA和BIOCLIM等在内的10种利用仅出现数据的建模方法对海洋底栖鱼类的栖息地进行模拟预测,其结果鼓励使用“仅出现”数据的方法,尤其是用MaxEnt来模拟底栖鱼类的栖息地;Reiss等[41]使用了包括GLM和MaxEnt等9种方法对20种底栖动物的分布进行模拟,结果表明MaxEnt的表现最为优异。剑鱼属于金枪鱼延绳钓渔业的兼捕渔获物,多数船队的渔捞日志数据质量较差,但由于在一定空间范围内记录出现点的精度要求很低,导致这类数据的可信度会大幅提高。本研究中模型预测能力的两个评价指标AUC值和模型遗漏率都支持MaxEnt能够为剑鱼这类非目标鱼种的栖息地模拟提供可靠工具。
本研究表明,海表温度是影响西印度洋剑鱼栖息地适宜性的主要因子,海表盐度和混合层深度是重要影响因子,而海面高度和叶绿素a浓度能够对剑鱼栖息地适宜性产生影响。这与北印度洋海域利用GAM和增强决策树(Boosted Regression Trees, BRT)对剑鱼分布和环境要素之间的关系研究结果相比,无论是环境因子的适宜范围还是剑鱼栖息地的偏好区域都很相似[42]。但是本研究发现,环境因子的重要性排序在干季和湿季会存在一定差异。
鱼类分布和洄游与温度密切相关[21],而剑鱼具有昼夜垂直移动的行为,白天通常在较深的水层活动,夜间移动到较浅水层[43]。因此,SST作为最容易获取的环境因子,在许多渔业研究中被采用[4-6, 9-10]。本研究结果表明,西印度洋区域剑鱼栖息地的适宜温度范围存在湿季和干季的差异,这与剑鱼进行产卵、索饵洄游和越冬洄游的时间选择和主动选择适宜温度的行为基本一致[44-45]。剑鱼的季节性纬向迁移也受到海温季节性变化的影响[8],干季由于季风洋流沿着阿拉伯海北部向西南方向与索马里暖流交汇,使赤道附近海域温度升高形成聚集区,且随着赤道逆流向东扩散[26];湿季受到南赤道环流影响,使剑鱼在30°S附近仍有分布区,同时索马里寒流向北运动,季风洋流在阿拉伯海海域向东运动[26],使剑鱼迁移至较温暖的阿拉伯海海域。
一些研究表明,海表盐度与混合层深度对剑鱼分布有重要影响[42],而印度洋西部和东部之间存在着盐度差[22],导致剑鱼的栖息地经向上分布有较为明显的差异。湿季在北印度洋,向东的西南季风将阿拉伯海的高盐水体输送至孟加拉湾[25];在南印度洋,湿季受到赤道与8°S之间的南赤道逆流影响,加上东南风,使高盐海水向赤道西部移动且向南北方向扩散[24]。海表盐度的变化与本研究中不同季节剑鱼栖息地的时空分布特征基本一致。混合层深度受到温度和盐度的影响[42],本研究中剑鱼栖息地主要分布在混合层相对较浅的区域。但同一年内湿季和干季之间存在较为明显的倾向差异,湿季剑鱼对混合层的深度倾向深于干季。受季风和环流影响,湿季西印度洋10°N以北海域存在一个较明显的深混合层碗状结构,迫使剑鱼向南选择栖息地;而干季剑鱼倾向于更浅的混合层,在赤道至20°S之间的海域成为主要栖息地[20]。强烈的季风触发表层和次表层营养物质的活化,从而导致叶绿素a浓度增加[42]。营养物质的增加,吸引了丰富的浮游生物。印度洋剑鱼主要捕食对象为中层鱼类和头足类,而头足类可以在浮游生物丰富的地方觅食[46]。因此,由于生理需求和饵料的可获得性,剑鱼倾向于在环境条件的适宜值范围内进行迁徙[42]
本研究利用MaxEnt对西印度洋剑鱼栖息地适宜性分布进行模拟预测,探讨了剑鱼在栖息地选择中的环境偏好。MaxEnt是找到在一定限制条件下熵最大的概率分布作为物种栖息地的最优分布[15]。所以限制条件的选取对模型的准确度必然存在影响。本文只选取了相关研究中常见的环境因子进行分析,建议在未来的研究中纳入更多的环境变量进行建模。另外,西印度洋作为世界上海洋生产力最高的区域之一,由于大气条件的季节性变化,初级生产力会受到海洋−大气交换的强烈影响[46],进而通过食物网的作用影响剑鱼饵料生物的分布,间接影响到剑鱼的迁移和空间分布,而目前对理解这些生物类群响应物理−生物过程的具体机制仍存在很大的局限性[47]。最后,印度洋是一个复杂的海区,同时具有复杂的气候模态[26],造就了不同的生物地理省[48],在后续研究中应该对研究区域进行更加细致的划分。
致谢:感谢中国远洋渔业数据中心和中国金枪鱼延绳钓渔业观察员为本研究提供西印度洋剑鱼渔获物的观察员数据,有力支持了剑鱼最大熵模型的建模过程,谨致谢忱!
  • 国家重点研发计划(2019YFD0901404)
  • 全球变化与海气相互作用专项(GASI-01-EIND-YD01aut/02aut)
  • 农业农村部远洋渔业国家观察员项目(17220255)
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2022年第44卷第10期
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doi: 10.12284/hyxb2022180
  • 接收时间:2022-04-14
  • 首发时间:2026-02-01
  • 出版时间:2022-10-01
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  • 收稿日期:2022-04-14
  • 修回日期:2022-06-07
基金
国家重点研发计划(2019YFD0901404)
全球变化与海气相互作用专项(GASI-01-EIND-YD01aut/02aut)
农业农村部远洋渔业国家观察员项目(17220255)
作者信息
    1.上海海洋大学 海洋科学学院,上海 201306
    2.国家远洋渔业工程技术研究中心,上海 201306
    3.大洋渔业资源可持续开发教育部重点实验室,上海 201306
    4.农业农村部大洋渔业资源环境科学观测实验站,上海 201306
    5.自然资源部第三海洋研究所,福建 厦门 361005

通讯作者:

王学昉(1983-),男,副教授,硕导,研究方向为海洋渔业资源监测与栖息地评估。E-mail:
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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

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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