Exhibit covariation characteristics in the habitat changes of Sardinops melanostictus and Scomber japonicus in the northwestern Pacific Ocean under ENSO event
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摘要: 远东拟沙丁鱼（Sardinops melanostictus）和日本鲭（Scomber japonicus）是西北太平洋海域重要的关联经济物种，探究二者栖息地变动的关联性有利于合理开发和管理渔业资源。本研究利用2017−2021年6−11月西北太平洋海域远东拟沙丁鱼和日本鲭的渔业数据，结合海表面温度、海表面高度和叶绿素a质量浓度3个关键环境变量分别构建不同权重的栖息地模型，并利用2021年的渔业数据进行验证。选取最优模型预测不同厄尔尼诺与南方涛动（El Niño-Southern Oscillation, ENSO）事件下远东拟沙丁鱼和日本鲭的最适栖息地分布，分析二者在不同ENSO事件下最适栖息地时空分布的差异性和同步性。结果表明：在不同ENSO事件下远东拟沙丁鱼适宜生境面积（高于15%）均高于日本鲭适宜生境面积（低于6%）；但远东拟沙丁鱼在拉尼娜事件下最适栖息地面积增长率高于厄尔尼诺事件，前者增长率为0.197，后者增长率为0.123，相反，日本鲭在拉尼娜事件下增长率低于厄尔尼诺事件，前者增长率为1.114，后者增长率为2.082；当远东拟沙丁鱼和日本鲭的分布位置接近时，会促进二者栖息地的适宜条件，当二者分布位置相距较远时会一定程度上抑制日本鲭栖息地面积的增加。远东拟沙丁鱼和日本鲭适宜面积在不同ENSO事件下协同变化特征可能与二者种间关系（竞争/捕食−被捕食）和西北太平洋海域海流分布情况有关。Abstract: The Sardinops melanostictus and the Scomber japonicus are important economic species in the northwestn Pacific Ocean, and exploring the correlation between their habitat changes is conducive to the rational development and management of fishery resources. This study utilizes fishery data of Sardinops melanostictus and Scomber japonicus in the northwestern Pacific Ocean from June to November between 2017 and 2021. By incorporating three key environmental variables, namely sea surface temperature, sea surface height, and chlorophyll a mass concentration, habitat models with different weights are constructed. The models are then validated using fishery data from 2021. The optimal models are selected to predict the most suitable habitat distribution of Sardinops melanostictus and Scomber japonicus under different El Niño-Southern Oscillation (ENSO) events. The study analyzes the differences and synchronicity in the spatial and temporal distribution of the most suitable habitat between the two species under different ENSO events. The results indicate that the suitable habitat area of Sardinops melanostictus (above 15%) was higher than that of Scomber japonicus (less than 6%) under different ENSO events; however, the growth rate of the most suitable habitat area for Sardinops melanostictus under La Niña events is higher than that of El Niño events. The former has a growth rate of 0.197 and the latter has a growth rate of 0.123, on the contrary, the growth rate of Scomber japonicus under the La Niña event is lower than that of El Niño event, the former has a growth rate of 1.114 and the latter has a growth rate of 2.082; additionally, when the distribution locations of Sardinops melanostictus and Scomber japonicus are close to each other, it promotes favorable conditions for their habitats. On the other hand, when the distribution locations are far apart, it somewhat inhibits the increase in the suitable habitat area for Scomber japonicus. The co-variation of suitable habitat areas for Sardinops melanostictus and Scomber japonicus under different ENSO events may be related to their interspecies relationship (competition/predation-prey) and the distribution of ocean currents in the northwestern Pacific Ocean.
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Key words:
- Sardinops melanostictus /
- Scomber japonicus /
- northwestern Pacific Ocean /
- El Niño /
- La Niña /
- habitat area
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图 1 各月环境变量拟合的远东拟沙丁鱼和日本鲭适宜性指数（SI）曲线
Fig. 1 Suitability index（SI）curve of each environmental variables for Sardinops melanostictus and Scomber japonicus in different months
图 2 2021年6−11月远东拟沙丁鱼（a）和日本鲭（b）捕捞努力量与栖息地适宜性指数（HSI）叠加分布
Fig. 2 The habitat suitability index (HSI) overlain with fishing efforts of Sardinops melanostictus (a) and Scomber japonicus (b) from June to November in 2021
图 3 不同ENSO事件下海表面温度、海表面高度、叶绿素a质量浓度时空分布
Fig. 3 Temporal and spatial distribution of sea surface temperature, sea surface height, and chlorophyll a mass concentration under different ENSO events
图 4 厄尔尼诺（2018年）和拉尼娜（2021年）事件下远东拟沙丁鱼（a）和日本鲭（b）6−11月适宜栖息地（HSI ≥ 0.6）时空分布
Fig. 4 Temporal and spatial distributions habitat (HSI ≥ 0.6) of Sardinops melanostictus (a) and Scomber japonicus (b) in El Niño event (2018) and La Niña event (2021) from June to November
图 5 厄尔尼诺（2018年）和拉尼娜（2021年）事件下远东拟沙丁鱼和日本鲭最适栖息地重心
Fig. 5 Optimal habitat center of gravity of Sardinops melanostictus and Scomber japonicus in El Niño event (2018) and La Niña event (2021)
图 6 厄尔尼诺（2018年）和拉尼娜（2021年）事件下6−11月远东拟沙丁鱼和日本鲭适宜栖息地（HSI ≥ 0.6）面积占比
Fig. 6 The proportion of suitable habitat area (HSI ≥ 0.6) of Sardinops melanostictus and Scomber japonicus in EI Niño event (2018) and La Niña event (2021) from June to November
图 7 不同ENSO气候事件下6−11月远东拟沙丁鱼和日本鲭最适栖息地中海表面温度、海表面高度、叶绿素a质量浓度均值
Fig. 7 Average values of sea surface temperature, sea surface height and chlorophyll a mass concentration in the optimal habitats of Sardinops melanostictus and Scomber japonicus from June to November under different ENSO climate events
表 1 渔业数据中建模和验证样本量
Tab. 1 Model and validate sample sizes in fishery data
物种 年份 样本量 6月 7月 8月 9月 10月 11月 远东拟沙丁鱼 2017 1 520 1 679 1 518 1 374 1 183 1 083 日本鲭 2017 1 616 1 686 1 538 1 311 1 095 1 078 远东拟沙丁鱼 2018 573 916 431 409 345 279 日本鲭 2018 1 681 1 653 1 068 1 047 721 471 远东拟沙丁鱼 2019 470 460 437 417 285 288 日本鲭 2019 510 543 518 439 339 298 远东拟沙丁鱼 2020 806 943 870 619 574 571 日本鲭 2020 636 773 643 534 731 647 远东拟沙丁鱼 2021 1 668 2 247 1 921 1 924 1 596 1 108 日本鲭 2021 886 810 316 374 633 497 注：“样本量”为2017−2021年各月作业天数的总和；2017−2020年的数据为建模数据，2021年的数据为验证数据。 
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表 2 不同关键环境变量的不同权重方案
Tab. 2 Different scenarios for the weights of the key environment variables
不同权重方案 海表面温度 海表面高度 叶绿素a质量浓度 方案一 0 1 0 方案二 0 0 1 方案三 0.1 0.8 0.1 方案四 0.1 0.1 0.8 方案五 0.25 0.5 0.25 方案六 0.25 0.25 0.5 方案七 0.5 0.25 0.25 方案八 0.333 0.333 0.333 方案九 0.8 0.1 0.1 方案十 1 0 0
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表 3 远东拟沙丁鱼和日本鲭SI模型拟合公式
Tab. 3 Fitting formula of SI models for Sardinops melanostictus and Scomber japonicus
月份 物种 SI 模型 R2 p 6月 远东拟沙丁鱼 SISST = exp[− 0.064 ×（XSST − 11.584）2] 0.795 <0.05 SIChl a = exp[− 33.686 ×（XChl a − 0.64）2] 0.966 <0.01 SISSH = exp[− 40.087 ×（XSSH − 0.237）2] 0.972 <0.01 日本鲭 SISST = exp[− 0.087 ×（XSST − 12.2）2] 0.795 <0.05 SIChl a = exp[− 141.335 ×（XChl a − 0.17）2] 0.902 <0.01 SISSH = exp[− 81.949 ×（XSSH − 0.228）2] 0.911 <0.01 7月 远东拟沙丁鱼 SISST = exp[− 0.069 ×（XSST − 16.429）2] 0.966 <0.01 SIChl a = exp[− 11.607 ×（XChl a − 0.267）2] 0.938 <0.01 SISSH = exp[− 54.679 ×（XSSH − 0.286）2] 0.976 <0.01 日本鲭 SISST = exp[− 0.069 ×（XSST − 15.658）2] 0.906 <0.01 SIChl a = exp[− 84.642 ×（XChl a − 0.195）2] 0.883 <0.05 SISSH = exp[− 66.081 ×（XSSH − 0.3）2] 0.961 <0.01 8月 远东拟沙丁鱼 SISST = exp[− 0.052 ×（XSST − 18.194）2] 0.987 <0.01 SIChl a = exp[− 204.505 ×（XChl a − 0.113）2] 0.890 <0.01 SISSH = exp[− 34.281 ×（XSSH − 0.303）2] 0.958 <0.01 日本鲭 SISST = exp[− 0.053 ×（XSST − 18.195）2] 0.957 <0.01 SIChl a = exp[− 51.625 ×（XChl a − 0.233）2] 0.926 <0.01 SISSH = exp[− 48.229 ×（XSSH − 0.297）2] 0.981 <0.01 9月 远东拟沙丁鱼 SISST = exp[− 0.053 ×（XSST − 15.85）2] 0.775 <0.05 SIChl a = exp[− 16.853 ×（XChl a − 0.096）2] 0.949 <0.01 SISSH = exp[− 87.937 ×（XSSH − 0.293）2] 0.918 <0.01 日本鲭 SISST = exp[− 0.027 ×（XSST − 17.05）2] 0.880 <0.01 SIChl a = exp[− 44.535 ×（XChl a − 0.294）2] 0.961 <0.01 SISSH = exp[− 30.036 ×（XSSH − 0.301）2] 0.919 <0.01 10月 远东拟沙丁鱼 SISST = exp[− 0.121 ×（XSST − 13.352）2] 0.875 <0.05 SIChl a = exp[− 83.825 ×（XChl a − 0.271）2] 0.981 <0.01 SISSH = exp[− 59.476 ×（XSSH − 0.318）2] 0.922 <0.01 日本鲭 SISST = exp[− 0.115 ×（XSST − 12.985）2] 0.944 <0.01 SIChl a = exp[− 45.493 ×（XChl a − 0.277）2] 0.833 <0.01 SISSH = exp[− 67.404 ×（XSSH − 0.313）2] 0.965 <0.01 11月 远东拟沙丁鱼 SISST = exp[− 0.158 ×（XSST − 10.131）2] 0.709 <0.05 SIChl a = exp[− 180.099 ×（XChl a − 0.392）2] 0.983 <0.01 SISSH = exp[− 65.196 ×（XSSH − 0.317）2] 0.857 <0.05 日本鲭 SISST = exp[− 0.136 ×（XSST − 10.817）2] 0.735 <0.01 SIChl a = exp[− 135.492 ×（XChl a − 0.19）2] 0.852 <0.01 SISSH = exp[− 54.85 ×（XSSH − 0.326）2] 0.878 <0.05
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表 4 厄尔尼诺（2018年）和拉尼娜（2021年）事件下6−11月远东拟沙丁鱼和日本鲭各月适宜栖息地（HSI≥0.6）面积增长率
Tab. 4 The growth rate of suitable habitat area (HSI ≥ 0.6) of Sardinops melanostictus and Scomber japonicus in El Niño event (2018) and La Niña event (2021) from June to November
物种 气候事件 7月 8月 9月 10月 11月 均值 远东拟沙丁鱼 El Niño 0.170 0.257 0.205 −0.004 −0.012 0.123 La Niña 0.984 0.210 0.080 −0.357 0.067 0.197 日本鲭 El Niño 9.041 0.291 1.240 −0.470 0.309 2.082 La Niña 0.485 0.052 2.826 −0.830 3.038 1.114
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