Landmark-based morphometric identification of two subpopulations of Larimichthys polyactis in Southern Yellow Sea and East China Sea
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摘要: 为比较不同地理群体的小黄鱼 (Larimichthys polyactis) 矢耳石形态特征，进一步明晰其种群划分问题，首次应用长于提取形态特征的地标点法探究了南黄海和东海小黄鱼群体的耳石形态学差异。2022年3—12月，自海州湾、吕泗、长江口和舟山渔场各采集30尾1龄小黄鱼，统一摘取其左矢耳石样本，在耳石外轮廓和耳石内听沟上分别提取了7个和4个地标点，进行地标点法分析。结果显示：1) 耳石听沟上的地标点8—11相对扭曲贡献率总计为91.88%，耳石外轮廓上的地标点1—7的总贡献率仅有8.12%，表明4个地理群体小黄鱼的耳石形态差异主要来源于耳石听沟；2) 主成分分析散点图显示，部分海州湾群体被显著的划分为1个类群，剩余部分海州湾渔场群体和其他3个渔场所有群体高度重叠聚集为另一个类群，表明南黄海海域可能存在两个小黄鱼种群，即南黄海-东海群和北黄海-渤海群，同时也表明海州湾渔场存在着小黄鱼混栖群体。该现象可能是由于小黄鱼早期鱼卵、仔幼鱼阶段被动扩散和后期主动洄游造成的生境差异，从而诱导其耳石形态表型性状特化。研究结果证实了耳石地标点法在鱼类种间识别的可行性，从耳石形态学角度可为中国近海小黄鱼种群划分提供参考。Abstract: To compare the otolith morphology of different Larimichthys polyactis populations, and to further clarify their population division, we applied the landmark method better at extracting morphological features for the first time, to explore the morphological differences of otoliths from L. polyactis populations in the Southern Yellow Sea and East China Sea. From March to December 2022, 30 individuals of 1-year old fish had been collected from each from Haizhou Bay (HZB), Lyusi (LS), Yangtze Estuary (YTE) and Zhoushan (ZS) fishing grounds. After extracting the left sagittal otoliths, we seclected even landmarks in the otolith contour and four landmarks in the sulcus acusticus for the landmark analysis. Results show that: 1) Among the 11 landmarks, those in the sulcus acusticus (Landmark 8−11) had main contribution (91.88%) in explaining the main source of morphological variation, while those in the otolith contour (Landmark 1−7) just had 8.12%, which reveals that the differences in otolith morphology among the four geographical groups mainly came from the auditory sulcus of the otolith. 2) According to the scatter plot of principal component analysis, part of the Haizhou Bay population was significantly divided into one group, while the remaining Haizhou Bay fishing grounds and all other three fishing grounds had highly overlapped and clustered into another group, indicating that there might be two L. polyactis populations in the Southern Yellow Sea area, namely the Southern Yellow Sea-East China Sea group and the Northern Yellow Sea-Bohai Sea group. This also indicates that there were mixed populations of L. polyactis in the Haizhou Bay fishing grounds. This may be explained by the long-term egg and larval dispersal process controlled by the current and active migratory routes during feeding and overwintering period. The results verified the potential of otolith morphology in delineating the populations of L. polyactis, providing valuable references for their spatiotemporal distribution.
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图 2 海州湾、吕泗、长江口和舟山渔场的小黄鱼左矢耳石内侧面11个地标点位置图
Figure 2. Positions of 11 landmarks on inner profile of left otolith morphology of L. polyactis sampled in Haizhou Bay (HZB), Lyusi (LS), Yangtze Estuary (YTE) and Zhoushan (ZS) fishing grounds
图 3 相对扭曲分数第1、第2主成分散点图 (主成分得分×102) 和网格变形图 (变异扩大10倍)
注：HZB. 海州湾渔场；LS. 吕泗渔场；YTE. 长江口渔场；ZS. 舟山渔场。图4同此。
Figure 3. Scatterplots of relative warp scores on 1st and 2nd (Principal component scores are enlarged 102 times) principal component and grid deformation of otolith (Variation are enlarged 10 times)
Note: HZB. Haizhou Bay fishing ground; LS. Lyusi fishing ground; YTE. Yangtze Estuary fishing ground; ZS. Zhoushan fishing ground. The same case in Fig. 4.
图 4 相对扭曲分数第1、第2、第3主成分3D散点图
Figure 4. 3D scatterplots of relative warp scores on 1st, 2nd and 3rd principal component
图 5 海州湾、吕泗、长江口和舟山渔场小黄矢耳石形态聚类分析结果
Figure 5. Dendrogram for hierarchical cluster analysis by otolith morphology of L. polyactis population in Haizhou Bay (HZB), Lvsi (LS), Yangtze Estuary (YTE) and Zhoushan (ZS) fishing grounds
表 1 4个渔场小黄鱼样本详细信息
Table 1. Detailed information of L. polyactis sampled from four fishing grounds
采样海域
Sampling sea area样本数
Number of samples采样时间
Sampling time体长
Body length/mm体质量
Body mass/g年龄
Age海州湾渔场
Haizhou Bay Fishing Ground, HZB30 2022年6—7月 108.30±4.54a 19.35±3.70 a 1+ 吕泗渔场
Lyusi Fishing Ground, LS30 2022年3—5月 107.90±6.65 a 19.20±3.85 a 1+ 长江口渔场
Yangtze Estuary Fishing Ground, YTE30 2022年8月 106.13±5.47 a 18.50±3.24 a 1+ 舟山渔场
Zhoushan Fishing Ground, ZS30 2022年11月 105.50±5.10 a 18.15±3.15 a 1+ 注：同列相同上标字母表示数据间无显著性差异 (P>0.05)。 Note: The same superscript letters within the same column indicate no significant differences among the data (P>0.05). 
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表 2 不同地标点的相对扭曲贡献率与定义
Table 2. Relative contribution and definitions of different landmarks
地标点
Landmark贡献率
Contribution rate/%定义
Definition地标点
Landmark贡献率
Contribution rate/%定义
Definition1 0.54 耳石背面的最宽点 7 0.54 听沟背面和前端之间突出的点 2 2.20 耳石背面和后端之间凹陷的点 8 4.71 听沟口端部和颈状部的靠耳石背面的交点 3 0.52 耳石后端的最长点 9 4.91 听沟口端部和颈状部的靠耳石腹面的交点 4 3.22 耳石腹面和后端之间凹陷的点 10 36.86 听沟尾端部和颈状部的靠耳石背面的交点 5 0.64 耳石腹面的最宽点 11 45.40 听沟尾端部和颈状部的靠耳石腹面的交点 6 0.45 耳石前端的最长点
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表 3 18个相对扭曲得分的特征值和贡献率
Table 3. Eigenvalues and contribution rates of 18 relative warps scores
主成分
Principal
component特征值
Eigenvalue贡献率
Contribution
rate/%累计贡献率
Cumulative contribution
rate/%1 2.49×10−2 89.93 89.93 2 7.23×10−4 2.21 92.13 3 6.43×10−4 1.96 94.10 4 4.04×10−4 1.23 95.33 5 3.38×10−4 1.03 96.36 6 2.08×10−4 0.64 97.00 7 1.93×10−4 0.59 97.59 8 1.83×10−4 0.56 98.15 9 1.33×10−4 0.41 98.55 10 1.16×10−4 0.35 98.91 11 8.70×10−5 0.26 99.17 12 5.30×10−5 0.16 99.33 13 5.00×10−5 0.15 99.49 14 4.80×10−5 0.15 99.64 15 4.00×10−5 0.12 99.76 16 3.30×10−5 0.10 99.86 17 2.80×10−5 0.09 99.95 18 1.80×10−5 0.05 100.00 总方差 Total variance 0.032 74
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