Speculation of migration routes of Larimichthys crocea in the East China Sea based on otolith microchemistry
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摘要: 近年来，东海频现对大黄鱼（Larimichthys crocea）大网捕获（吨级规模以上），引起了对衰退中的野生大黄鱼资源保护的担忧。为查明东海野生大黄鱼洄游规律，本文根据2020−2022年在浙江近海产卵场和外海越冬场采集的不同批次野生大黄鱼样本，利用激光剥蚀电感耦合等离子体质谱法（LA-ICP-MS）测定耳石微化学元素，结合聚类分析和主成分分析（PCA），对不同大黄鱼群体进行了划分，结合耳石的年轮特征，研究大黄鱼群体的栖息生境，推测大黄鱼洄游路线。结果表明，Ba/Ca
$\leqslant$ 0.004 2时可以判定为海洋栖息生境，Ba/Ca$\geqslant$ 0 .008 1时可以判定为河口栖息生境，0.004 2 < Ba/Ca< 0.008 1时可以判定为混合栖息生境。根据该阈值范围，判定本次采样的大黄鱼的5种生境类型：（1）近海产卵，近海混合水域短暂栖息类型（占22.2%）；（2）近海产卵，阶段性往复混合生境栖息类型（占15.6%）；（3）近海产卵，河口水域育幼索饵，外海越冬栖息类型（31.1%）；（4）河口产卵、混合水域育幼，外海越冬栖息类型（6.7%）；（5）近海混合水域短暂产卵，绝大部分时间海洋生境栖息类型（24.4%）。对所有样本的Ba/Ca核心、峰值和边缘值进行PCA，5种栖息类型的样本均匀分布在第一轴的两侧，大部分时间都在海洋生境中的G1、G2和G5，以及受陆源影响较大的G3和G4。同时，PCA结果同时也显示，5种类型样本均会近同步地在近海岛礁处出现，这表明外海、近海和河口的群体由于洄游等因素的影响会出现混合，说明了舟山岛礁水域对大黄鱼产卵育幼的重要性。本研究为东海大黄鱼近海产卵场、索饵场和外海越冬场之间洄游履历，以及大黄鱼洄游路线推测提供一定依据。Abstract: In recent years, large yellow croaker (LYC) has been caught with high abundance in the East China Sea (ECS), which raises concerns about the conservation of LYC resources. We collected LYC samples from spawning grounds and wintering grounds in the offshore Zhejiang from 2020 to 2022. In order to find out the migration route of LYC in the ECS, we made use of otolith microchemical elements by LA-ICP-MS analysis, combined with the clustering analysis and PCA of trace elements such as Ba/Ca, to classify different LYC groups. With the age characteristics of otolith, the habitat patterns of LYC group was studied, and the updated migration route of LYC was speculated. The results showed that groups with Ba/Ca$\leqslant$ 0.004 2 was inhabited in the marine region. That with Ba/Ca$\geqslant$ 0.008 1 was inhabited in the estuary region, and that with 0.004 2 < Ba/Ca < 0.008 1 was inhabited in the mixed (estuary and oceanic) habitat. The five group types of LYC were: (1) offshore spawning, short-term mixed waters inhabited (accounting for 22.2%); (2) offshore spawning, periodically back and forth with mixed water inhabited (accounting for 15.6%); (3) offshore spawning, feeding and nursering in estuary waters, and wintering inhabited in the open sea (31.1%); (4) estuary spawning, nursering in the mixed waters, overwintering inhabited in the open sea (6.7%); (5) shortly spawning in the mixed waters, most of the time inhabited in the open ocean (24.4%). PCA was performed with Ba/Ca core, peak and edge values of all samples, and the results showed that samples from five groups were evenly distributed on both sides of the first axis: G1, G2 and G5 groups, which spent most of their time in the ocean, and G3 and G4 groups, which were heavily influenced by terrestrial sources. At the same time, the PCA results also showed that all the samples appeared nearly synchronously near the island or reefs, indicating that populations in the open sea, offshore and estuary waters may mix due to factors such as migration, indicating the importance of Zhoushan offshore waters for LYC nursering and spawning. This study provides a certain basis for the migration history of the LYC in the ECS between its coastal spawning, feeding grounds and its overwintering grounds in the open sea, as well as for the speculation of its migration routes. -
图 2 大黄鱼样品的采集站位
近岸和外海样本是指渔获来源地，右图上的数字指样品编号，根据测定的批次和时间顺序共同决定
Fig. 2 The sampling stations of Larimichthys crocea
Nearshore and offshore samples indicates sampling place, numbers in right figure indicates the sampling code, decided by the sequence of processing and sampling date
图 4 样本聚类图（数字表示样品编号）
Fig. 4 Hierarchical clustering of samples (numbers indicate sample code)
图 6 G3所有样本耳石平均Ba/Ca（a）及突变概率（b）
黑圈状折线为G3从耳石核心至边缘的Ba/Ca分布，绿色线为概率，红色、蓝色和绿色阴影区分别表示0峰值阶段，前后第一次突变，前后第二次突变对应的折线
Fig. 6 Average of Ba/Ca (a) and its trend change probability (b) in G3 otolith samples
Black circle line indicate Ba/Ca curve from otolith core to edge in G3, green line indicate probability, red, blue and green shade indicates peak, first trend change point lines, and second change point lines
图 7 大黄鱼5个群体的Ba/Ca值变化
蓝色、棕色和黄色分别表示海洋、混合和河口生境
Fig. 7 Changes of Ba/Ca values in five groups of Larimichthys crocea
Blue, brown and yellow indicates oceanic, mixed water and estuary conditions
图 8 G1−G5 5个代表性大黄鱼耳石的Ba/Ca轨迹
灰色竖状阴影区为冬轮，非阴影区为夏轮
Fig. 8 Ba/Ca trajectories corresponding to the otoliths of Group 1 to Group 5 of Larimichthys crocea
Vertical grey shaded area indicates winter growth ring, non-shaded indicates summer growth ring
图 9 45个样本的Ba/Ca核心区、中间区和边缘区的PCA（a）和代表性样本Ba/Ca的PCA与季节、地理变化（b）
Fig. 9 PCA of Ba/Ca core, intermediate and edge value (a) and PCA of the selected samples and its seasonal/geographical variation (b)
图 10 大黄鱼洄游路线推测
橘色点表示岛礁外侧产卵群体；黄色点表示岛礁区内近河口群体；绿色点表示混合群体
Fig. 10 The speculative migratory route of Larimichthys crocea
Orange dots indicate offshore spawning groups; yellow dots indicate nearshore-estuary groups; green dots indicate mixed groups
表 1 大黄鱼的样品采样信息
Tab. 1 Sampling details of Larimichthys crocea in the present study
年份 季节 样本数量 样本编号 体长范围
/cm体质量范围
/g2020 秋季 6* 1，2，23～26 29.7～33.4 334.7～437.4 2021 春季 1 46 30.5 427 2021 秋季 9 27～35 13.9～16.9 48.1～74.3 2022 春季 8* 19～22，42～44，48 13.3～27.0 30.5～109.1 2022 秋季 14 5～18 9.5～41.0 16.3～646.0 2022 冬季 7* 3～4，36～38，47，49 25.0～44.8 234.8～1144.3 注：*代表部分样本无体长、体质量数据。 
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表 2 东海大黄鱼耳石Mg、Mn、Sr、Ba 4种元素的变异程度
Tab. 2 Degree of variations of Mg, Mn, Sr, Ba in the otoliths of Larimichthys crocea in the East China Sea
指标 均值/(mmol·mol−1) 标准差/(mmol·mol−1) 最小值
/(mmol·mol−1)中位数
/(mmol·mol−1)最大值
/(mmol·mol−1)变异系数
/%Sr/Ca 2.36 0.56 0.67 2.28 7.28 23.50 Ba/Ca 4.76 × 10−3 0.01 1.06 × 10−4 3.4 × 10−3 0.09 112.05 Mg/Ca 0.08 0.06 0.01 0.06 3.46 86.42 Mn/Ca 0.01 0.01 4.49 × 10−5 0.01 0.15 75.72
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表 3 4个组群Ba/Ca的变异程度
Tab. 3 Variation degree of Ba/Ca in four groups
分组 均值/(mmol mol−1) 标准差/(mmol mol−1) 变异系数/% G1 3.85 × 10−3 1.88 × 10−3 48.12 G2 3.83 × 10−3 1.6 × 10−3 42.15 G3 7.4 × 10−3 6.38 × 10−3 84.88 G4 3.2 × 10−3 2.25 × 10−3 55.31 G5 3.36 × 10−3 1.21 × 10−3 38.33
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