A comparative analysis of green tides in the North Yellow Sea and South Yellow Sea: perspectives from spatiotemporal distribution and genetic sequencing
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摘要: 2008年以来南黄海持续遭遇浒苔绿潮灾害，2009年以来北黄海也频发大型藻类堆积的现象。大规模绿潮暴发严重破坏海洋生态，对近海生态平衡和经济结构造成威胁，然而对南北黄海绿潮的暴发差异研究甚少。本文基于多源遥感数据、无人机监测和分子生物学方法，从时空分布格局和遗传基础两个互补维度，系统性地对比研究了南北黄海浒苔的差异性。研究结果显示：（1）暴发时间和暴发规模显著性差异。南黄海浒苔群落最早出现于5月份，最大覆盖面积通常出现在6月中下旬，至7−8月间逐渐消失，整个生命周期持续约为50～90 d。北黄海浒苔为本土生浒苔，6月底至7月初出现在烟台近海，8月份逐渐消失，较南黄海浒苔覆盖面积小、持续时间短；近年南黄海浒苔暴发面积均大于200 km2，北黄海浒苔暴发面积大致保持在
2000 m2以内，显著小于南黄海海域。（2）2018−2024年南黄海浒苔的迁移路径可归纳为北向漂移型和先北后南型两类。在空间分布格局上，南黄海浒苔藻华呈现大规模连片聚集特征，而北黄海则表现为局部零星分布。南黄海浒苔起源于苏北浅滩，借季风与海流北迁至山东半岛南部聚集；北黄海浒苔主要集中在烟台市开发区金沙滩、莱山区逛荡河入海口等区域，无大范围漂移；（3）ITS基因测序表明，烟台近岸北黄海浒苔属于一种新的遗传品系（ITS JST3），其浅绿色短小丛生状形态、发育不完善气囊结构显著区别于南黄海浒苔。本研究从宏观时空格局与微观遗传基础相结合的维度，系统证实了南北黄海绿潮为驱动机制不同的独立生态事件，对绿潮的分区精准防控具有重要指导意义。Abstract: Since 2008, the South Yellow Sea has been persistently affected by green tide disasters caused by Ulva prolifera, while since 2009, the North Yellow Sea has frequently experienced large-scale macroalgal accumulations. The massive outbreaks of green tides severely damage marine ecosystems and threaten coastal ecological balance and economic structures. However, research on the differences in green tide outbreaks between the South Yellow Sea and North Yellow Sea remains limited. This study systematically compares the differences in Ulva prolifera between the two regions from two complementary dimensions—spatiotemporal distribution patterns and genetic basis—using multi-source remote sensing data, UAV monitoring, and molecular biology methods. The results indicate: (1) Significant differences in outbreak timing and scale. Ulva prolifera in the South Yellow Sea first appears in May, reaches its maximum coverage in mid-to-late June, and gradually disappears between July and August, with an entire life cycle lasting approximately 50–90 d. In contrast, Ulva prolifera in the North Yellow Sea is locally sourced, appearing in the coastal waters of Yantai from late June to early July and gradually disappearing in August. It exhibits smaller coverage areas and shorter duration compared to the South Yellow Sea. In recent years, the outbreak area in the South Yellow Sea has consistently exceeded 200 km2, while that in the North Yellow Sea has remained within2000 m2, significantly smaller. (2) From 2018 to 2024, the migration paths of Ulva prolifera in the South Yellow Sea can be categorized into two types: northward drifting and northward-then-southward drifting. Spatially, Ulva prolifera blooms in the South Yellow Sea exhibit large-scale, continuous aggregation, whereas those in the North Yellow Sea are characterized by localized, scattered distributions. Ulva prolifera in the South Yellow Sea originates from the Subei Shoal, drifting northward with monsoon winds and ocean currents to accumulate along the southern coast of the Shandong Peninsula. In the North Yellow Sea, Ulva prolifera is mainly concentrated in areas such as the Jinshatan Beach in Yantai Development Zone and the estuary of the Guangdang River in Laishan District, with no large-scale drifting observed. (3) ITS gene sequencing reveals that Ulva prolifera in the North Yellow Sea along the Yantai coast belongs to a novel genetic strain (ITS JST3), which is distinctly different from the South Yellow Sea strain in terms of its light green, short, clustered morphology and underdeveloped air sac structures. By integrating macroscopic spatiotemporal patterns with microscopic genetic foundations, this study systematically demonstrates that the green tides in the South and North Yellow Sea are independent ecological events driven by different mechanisms, providing important insights for targeted regional prevention and control of green tides.-
Key words:
- Ulva prolifera /
- spatiotemporal characteristics /
- Yellow Sea /
- ITS gene sequences /
- divergence
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图 1 研究区位置分别为烟台市（a）、海阳市（b）、日照市（c）无人机实地观测影像
Fig. 1 Location of study area are drone field observation images from Yantai City (a), Haiyang City (b), and Rizhao City (c), respectively
图 2 浒苔、植被、海水的光谱反射率曲线
Fig. 2 Spectral reflectance curves of Ulva prolifera, terrestrial vegetation, and seawater
图 3 南黄海浒苔时空分布（以2024年为例）
图中白色区域表示云覆盖，该区域无法进行有效光学观测
Fig. 3 Spatiotemporal distribution of Ulva prolifera in the South Yellow Sea (2024 representative case)
Areas with white diagonal lines indicate cloud cover, where valid optical observation was not possible
图 4 2024年6月26日烟台开发区金沙滩（a、b）2024年7月5日烟台市莱山区（c、d）
Fig. 4 Golden Beach, Yantai Development Zone (26 June 2024) (a, b) Laishan District, Yantai City (5 July 2024) (c, d)
图 5 2018−2024年南黄海浒苔的漂移路径
Fig. 5 Drift trajectories of Ulva prolifera in the South Yellow Sea (2018–2024)
图 6 2018−2024年南北黄海浒苔最大覆盖面积趋势图
Fig. 6 Interannual variation in maximum coverage area of Ulva prolifera in North Yellow Sea and South Yellow Sea (2018–2024)
图 7 南北黄海浒苔ITS 系统发育树
▲表示北黄海浒苔优势种
Fig. 7 Phylogenetic tree based on ITS sequences of Ulva prolifera from North Yellow Sea and South Yellow Sea
▲ denotes dominant species in North Yellow Sea
表 1 2018−2024年遥感影像成像日期
Tab. 1 Acquisition dates of remote sensing images from 2018 to 2024
年份 GF-1数据 HJ-1/2数据 Sentinel-2数据 HY-1C数据 2018 7月27日 − 6月8日，6月11日，6月21日，6月29日，7月3日，7月6日，7月16日，
7月21日，8月2日，8月7日，8月10日− 2019 7月3日，7月21日 − 5月9日，5月20日，5月29日，6月2日，6月3日，6月10日，6月18日，
6月23日，7月1日，7月8日，7月10日，7月26日，8月5日，8月7日− 2020 7月25日，8月4日 − 5月18日，5月23日，5月26日，6月1日，6月4日，6月21日，6月25日，
7月1日，7月10日，7月15日− 2021 7月10日，7月25日 − 5月21日，5月28日，6月6日，6月20日，6月23日，7月2日，7月14日，
7月18日，7月21日，8月1日，8月11日，8月14日，8月16日，8月19日− 2022 7月25日 7月16日 5月22日，5月25日，5月30日，6月15日，6月18日，6月25日 − 2023 6月15日 7月5日，7月19日，
7月23日− 5月14日，5月23日，6月8日，6月22日 2024 5月16日，6月6日，6月10日，
6月18日，6月23日，7月21日5月28日，6月1日 − 6月25日，8月3日 注：“−”表示无成像。 
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