Diversity, distribution and environmental correlation of harmful dinoflagellate cysts in the Bohai Sea and the North Yellow Sea in spring

doi:10.17520/biods.2025426

Abstract

Abstract:

Aims: The frequency of harmful algal blooms (HABs) in the Bohai Sea and the North Yellow Sea has been increasing continuously in the context of coastal eutrophication and climate change. Dinoflagellate cysts are considered a potential seed bank and play a key role in the occurrence of HABs. This study aimed to investigate the diversity, distribution patterns, and environmental factors of harmful dinoflagellate cysts in the Bohai Sea and the North Yellow Sea, and to assess the potential risk of HABs in these regions.

Methods: A DNA metabarcoding approach was applied, targeting the small subunit ribosomal (18S V4) and internal transcribed spacer (ITS1) to investigate the diversity and distribution of harmful dinoflagellate cyst assemblages in surface sediments from the Bohai Sea and the North Yellow Sea in May 2023. Canonical correspondence analysis (CCA) was used to assess the correlations between cyst assemblages and environmental factors.

Results: The environmental DNA (eDNA) information revealed 38 potentially harmful dinoflagellate cyst species, including 5 species not previously known to produce dinoflagellate cysts, and 8 newly recorded dinoflagellate cysts species of the Bohai Sea and the North Yellow Sea. Among them, Azadinium spinosum, Gymnodinium nolleriand Alexandrium fragae were detected for the first time in sediment samples from the coastal waters of China. Alpha diversity analysis revealed a clear decreasing trend in species richness from the Bohai Sea, through the Bohai Strait, to the central North Yellow Sea, and this trend was positively correlated with the water temperature gradient. CCA result indicated that the abundance and distribution of dominant harmful dinoflagellate cysts in the two seas were influenced to varying degrees by environmental factors (temperature, salinity, water depth, dissolved inorganic nitrogen (DIN), dissolved inorganic phosphorus (DIP) and N/P), with temperature being the main explanatory variable. Temperature, in combination with the N/P and DIN, affected and regulated the abundance and distribution of most coastal-estuarine species, resulting in higher detection rates and/or abundances in the warm and eutrophic southwestern Bohai Sea. Low temperature, high salinity, and high phosphorus levels were key factors contributing to the dominance of Alexandrium catenella cysts in the North Yellow Sea, particularly around the central cold-water mass.

Conclusion: This study provides crucial information on the diversity and distribution patterns of harmful dinoflagellate cysts in the whole Bohai Sea and the North Yellow Sea in spring. The southwestern Bohai Sea remains a high-risk area for HABs due to both established toxic and harmful species and emerging potential risks, such as HABs-forming, Lepidodinium chlorophorum, whereas the North Yellow Sea faces elevated risks from the widespread high relative abundance of Alexandrium catenella. These findings demonstrate that molecular techniques are a powerful tool for monitoring harmful dinoflagellate cysts, underscoring the need for their use in early warning of HABs and for protecting marine aquaculture from blooms.

Key words: harmful dinoflagellate cysts, DNA metabarcoding, diversity, distribution, environmental factors

Zhicheng Kang, Chunlei Gao, Jianing Guo, Fanping Meng, Zongling Wang. Diversity, distribution and environmental correlation of harmful dinoflagellate cysts in the Bohai Sea and the North Yellow Sea in spring[J]. Biodiv Sci, 2026, 34(3): 25426.

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URL: https://www.biodiversity-science.net/EN/10.17520/biods.2025426

https://www.biodiversity-science.net/EN/Y2026/V34/I3/25426

Figures/Tables 13

Fig. 1 Sampling stations in the Bohai Sea and the North Yellow Sea, China

Table 1 The primers used for high-throughput sequencing in this study

扩增区域 Target region	扩增片段长度 Fragment length	引物名称 Primers	引物序列(5′-3′) Sequences of primers (5′-3′)	文献 References
18S V4	370 bp	TAReuk454FWD1	CCAGCA(G/C)C(C/T)GCGGTAATTCC	Stoeck et al., 2010
		TAReukREV3	ACTTTCGTTCTTGAT(C/T)(A/G)A	Stoeck et al., 2010
ITS1 (5.8S)	260 bp	PRIMER B-reverse	TAGGTGAACCTGCAGAAGGAT	Medlin et al., 1988
		ITS300R	CACGGAA(G/T)TTCTGCA(A/G)TTCACAATG	Scholin et al., 1994

Table 2 PCR amplification reaction system for high-throughput sequencing

组分 Components	体积 Volume (μL)	浓度 Concentration
DNA模板 DNA template	2	含量 Amount < 200 ng
正向引物 Forward primer	1	10 μM
反向引物 Reverse primer	1	10 μM
dNTP Mix	4	2.5 mM
Ex Taq DNA聚合酶 Ex Taq DNA polymerase (Takara)	0.3	50 U/μL
10× Ex Taq buffer	5
双蒸水 ddH₂O	36.7
总体积 Total volume	50

Fig. 2 Distribution of α diversity indices of harmful dinoflagellate cyst in the Bohai Sea and the North Yellow Sea, China

Fig. 3 Result of principal coordinate analysis (PCoA) of harmful dinoflagellate cyst assemblages in the Bohai Sea and the North Yellow Sea, China. A, Stations in the sea area near the mouth of Bohai Bay; B, Stations along the coast of the North Yellow Sea and in the Bohai Sea area outside the mouth of Bohai Bay; C, Stations in the central North Yellow Sea.

Fig. 4 Relative abundance of 12 dominant genera (relative abundance > 5%) at each sampling station

Fig. 5 Distribution of relative abundance of cysts in the dominant genus (relative abundance > 5%)

Fig. 6 Relative abundance of dominant species (relative abundance > 10%) and sub-dominant species (relative abundance 5%-10%) at each station. Black, Dominant species; Blue, Sub-dominant species.

Fig. 7 Distribution of relative abundance of dominant and sub-dominant cysts in species level

Fig. 8 Canonical Correspondence Analysis (CCA) performed on the relative abundance of dominant and sub-dominant harmful dinoflagellate cysts and in-situ environmental parameters. A. aff, Alexandrium affine; A. and, Alexandrium andersonii; A. cat, Alexandrium catenella; A. leei: Alexandrium leei; A. min, Alexandrium minutum; A. ost, Alexandrium ostenfeldii; A. pseu, Alexandrium pseudogonyaulax; Aza. spi, Azadinium spinosum; B. bre, Biecheleria brevisulcata; G. spi, Gonyaulax spinifera; Gym. imp, Gymnodinium impudicum; H. ste, Heterocapsa steinii; L. chl, Lepidodinium chlorophorum; Pen. dal, Pentapharsodinium dalei; Po. har, Polykrikos hartmannii; Po. kof, Polykrikos kofoidii; Pro. exc, Protoperidinium excentricum; Pyr. Ste, Pyrophacus steinii; S. acu, Scrippsiella acuminata. DIN, Dissolved inorganic nitrogen; DIP, Dissolved inorganic phosphorus.

Table 3 The independent explanatory rate and significance of each environmental factors

环境因素 Environmental factors	解释度 Explaination (%)	P
温度 Temperature	15.93	0.002
水深 Water depth	13.81	0.002
盐度 Salinity	12.44	0.002
可溶性无机氮 Dissolved inorganic nitrogen	3.72	0.004
氮磷比 N/P	3.34	0.008
可溶性无机磷 Dissolved inorganic phosphorus	2.61	0.016

Fig. 9 Distribution of in-situ environmental factors. (a) Temperature; (b) Salinity; (c) Dissolved inorganic phosphorus (DIP); (d) Dissolved inorganic nitrogen (DIN); (e) N/P.

Fig. 10 Distribution of eDNA of new recorded harmful algal blooms dinoflagellate Lepidodinium chlorophorum in the surface sediments of the Bohai Sea and the North Yellow Sea

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