中国沿海洛氏角毛藻复合群的多样性组成及地理分布

doi:10.17520/biods.2018261

生物多样性 ›› 2019, Vol. 27 ›› Issue (2): 149-158. DOI: 10.17520/biods.2018261

中国沿海洛氏角毛藻复合群的多样性组成及地理分布

陈作艺^1,²,许晓静¹,朱素英¹,翟梦怡¹,李扬^1,^*()

1 华南师范大学生命科学学院, 广州市亚热带生物多样性与环境生物监测重点实验室, 广东省水产健康安全养殖重点实验室, 广州 510631
2 河北省地矿局第八地质大队, 河北秦皇岛 066001

收稿日期:2018-09-28 接受日期:2019-01-27 出版日期:2019-02-20 发布日期:2019-04-16
通讯作者: 李扬
基金资助:
国家自然科学基金(31570205);科技基础资源调查专项(2018FY100200)

Species diversity and geographical distribution of the Chaetoceros lorenzianus complex along the coast of China

Chen Zuoyi^1,²,Xu Xiaojing¹,Zhu Suying¹,Zhai Mengyi¹,Li Yang^1,^*()

1 Guangzhou Key Laboratory of Subtropical Biodiversity and Biomonitoring, Guangdong Provincial Key Laboratory of Healthy and Safe Aquaculture, School of Life Sciences, South China Normal University, Guangzhou 510631
2 The Eighth Geological Brigade, Hebei Geological Prospecting Bureau, Qinhuangdao, Hebei 066001

Received:2018-09-28 Accepted:2019-01-27 Online:2019-02-20 Published:2019-04-16
Contact: Li Yang

摘要/Abstract

摘要：

洛氏角毛藻复合群(Chaetoceros lorenzianus complex)指具有与洛氏角毛藻相似形态学特征的物种集合, 它们广泛分布于全球近岸水域。近年国际上关于该复合群的分类学研究取得新进展, 而我国相关研究仍较为滞后。为了弄清我国沿海洛氏角毛藻复合群的物种多样性, 明确物种信息, 厘清种间界限, 为相关研究提供准确的物种鉴定依据, 本研究陆续在中国沿海建立了该复合群的332个单克隆培养株系, 利用光学显微镜、扫描电镜和透射电镜进行了较为详尽的形态学研究, 基于核糖体大亚基编码基因D1-D3区序列, 构建了分子系统学关系。结果表明其形态聚类与分子系统学结论相一致, 显示我国洛氏角毛藻复合群具有较高的物种多样性, 共鉴定到5个物种, 分别是并基角毛藻(C. decipiens)、优美角毛藻(C. elegans)、平孢角毛藻(C. laevisporus)、曼纳角毛藻(C. mannaii)和稀树角毛藻(C. pauciramosus)。研究表明传统认知的光镜下特征, 如群体特征、角毛走势等易变化, 其分类学价值需谨慎应用。角毛的超微结构, 如角毛孔纹的形状、大小、密度等是有效的种间区别特征, 休眠孢子亦是重要的物种识别依据。并基角毛藻和平孢角毛藻在我国沿岸的分布范围最为广泛, 而稀树角毛藻的分布较为有限。

关键词: 洛氏角毛藻复合群, 物种多样性, 形态学, 分子系统学, 休眠孢子, 中国沿海

Abstract

The Chaetoceros lorenzianus complex is composed of several planktonic diatom species that share a similar morphology with C. lorenzianus Grunow. The complex has been recorded frequently in coastal waters across the world. Recently, new taxonomic information has enriched the known species diversity of the complex from 3 to 7 taxa. However, the identities of many species comprising the complex in China is still unclear. To clarify the species diversity and provide solid identification criteria for further studies, 332 monoclonal strains belonging to the C. lorenzianus complex were collected from the coast of China. With light microscopy and scanning and transmission electron microscopy, the morphologies of vegetative cells and resting spores were observed. Hypervariable D1-D3 regions of nuclear large subunit ribosomal encoded genes were amplified to construct the phylogenetic relationship. Morphological clustering was consistent with molecular systematics, which indicated that a total of 5 species are present within the C. lorenzianus complex along the coast of China; C. decipiens, C. elegans, C. laevisporus, C. mannaii and C. pauciramosus. The ultrastructure on setae, such as the shape, size and density of the setae pores, can be used as distinguishing features among allied taxa. The morphology of the resting spores also aid in accurate identification. The previous reports identifying the taxa within the C. lorenzianus complex in China need to be reexamined. Chaetoceros decipiens and C. laevisporus are the most widespread taxa along the coast of China, whereas C. pauciramosus has the narrowest distribution, having only been recorded in Taishan and Zhuhai of the Guangdong Province.

Key words: Chaetoceros lorenzianus complex, species diversity, morphology, molecular phylogeny, resting spore, coast of China

陈作艺, 许晓静, 朱素英, 翟梦怡, 李扬 (2019) 中国沿海洛氏角毛藻复合群的多样性组成及地理分布. 生物多样性, 27, 149-158. DOI: 10.17520/biods.2018261.

Chen Zuoyi, Xu Xiaojing, Zhu Suying, Zhai Mengyi, Li Yang (2019) Species diversity and geographical distribution of the Chaetoceros lorenzianus complex along the coast of China. Biodiversity Science, 27, 149-158. DOI: 10.17520/biods.2018261.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2018261

https://www.biodiversity-science.net/CN/Y2019/V27/I2/149

图/表 9

图1 采样站位及中国沿海洛氏角毛藻复合群物种地理分布图。〇: 采样站位; A: 渤海; B: 山东半岛; C: 浙江沿岸; D: 台湾海峡; E: 广东沿岸; F: 环海南岛。

Fig. 1 Sampling sites and geographical distribution of the Chaetoceros lorenzianus complex along the coast of China. 〇, Sampling sites; A, Bohai Sea; B, Shandong coast; C, Zhejiang coast; D, Taiwan Strait; E, Guangdong coast; F, Hainan coast.

图2 并基角毛藻的光学显微镜(LM) (A, E)、扫描电镜(SEM) (B-D, F, G)和透射电镜(TEM) (H-J)示意图。A-B: 细胞链宽环面观, 分别显示链中角毛无并行融合(A, 箭头)及有并行融合(B, 箭头); C: 壳面观; D: 链端壳面, 示U型端角毛及唇形突(箭头); E-G: 角毛结构; H: 链端壳面, 示唇形突(箭型); I: 壳套; J: 环带。标尺 = 20 μm (A), 10 μm (B, D, E), 6 μm (C), 5 μm (H), 2 μm (F, G, J), 1 μm (I)。

Fig. 2 Morphology of Chaetoceros decipiens under light microscopy (LM) (A, E), scanning electron microscopy (SEM) (B-D, F, G) and transmission electron microscopy (TEM) (H-J). A-B, Broad girdle views showing fusing sibling setae base present (B, arrowheads) and not (A, arrowheads); C, Internal view of intercalary valve; D, Terminal valve with rimoportula (arrowhead) and U-shaped terminal setae; E-G, Structure of setae; H, Terminal valve with rimoportula (arrow); I, Mantle; J, Girdle bands. Scale bars, 20 μm (A), 10 μm (B, D, E), 6 μm (C), 5 μm (H), 2 μm (F, G, J), 1 μm (I).

图3 优美角毛藻的光学显微镜(LM) (A-B)、扫描电镜(SEM) (C-D, I-J)和透射电镜(TEM) (E-H)示意图。A: 链状群体; B: 休眠孢子位于母细胞链中; C: 链中壳面, 示硅质翼(箭头); D: 释放的成熟休眠孢子; E: 端壳面, 示唇形突(箭头)和硅质脊(箭型); F: 壳套; G: 链端壳面, 示中央环纹(箭型); H: 环带; I-J: 角毛结构. 标尺 = 20 μm (A, B), 6 μm (D), 4 μm (C), 2 μm (E, F, G, H, I, J)。

Fig. 3 Morphology of Chaetoceros elegans under light microscopy (LM) (A-B), scanning electron microscopy (SEM) (C-D, I-J) and transmission electron microscopy (TEM) (E-H). A, Chain in broad girdle view; B, Resting spores within the mother cells of a chain; C, Sibling intercalary valves showing overlapping silica ear-like structures (arrowhead); D, Released resting spore. E, Terminal valve with external process of rimoportula (arrowhead) and fringe (arrow); F, Parallel rows of poroids on the mantle; G, Central annulus (arrow) on terminal valve; H, Girdle bands; I and J, Seta structure. Scale bars, 20 μm (A, B), 6 μm (D), 4 μm (C), 2 μm (E, F, G, H, I, J).

图4 平孢角毛藻的光学显微镜(LM) (A-B)、扫描电镜(SEM) (C-E, I-J)和透射电镜(TEM) (F-H)示意图。A: 链状群体; B: 休眠孢子位于母细胞链中; C: 链中壳面, 示硅质翼(宽箭型), 硅质脊(箭头)及壳套基部凹槽(窄箭型); D: 释放的成熟休眠孢子; E: 链端壳面, 示唇形突(箭头)及硅质肋纹(箭型); F: 壳套; G: 链中壳面, 示中央环纹(箭型); H: 环带; I, J: 角毛结构。标尺 = 50 μm (A), 20 μm (B), 4 μm (C, E), 2 μm (D, F, G, H, I, J)。

Fig. 4 Morphology of Chaetoceros laevisporus under light microscopy (LM) (A-B), scanning electron microscopy (SEM) (C-E, I-J) and transmission electron microscopy (TEM) (F-H). A, Chain in broad girdle view; B, Resting spores within the mother cells of a chain; C, Sibling intercalary valves showing overlapping silica wings (broad arrow), silica ridges (arrowhead) and furrow above the basal ring of mantle (arrow); D, A released resting spore; E, Terminal valve with central processes (arrowhead) and silica rib (arrows); F, Mantle; G, Intercalary valve with central annulus (arrow); H, Girdle bands; I and J, Setae structure. Scale bars, 50 μm (A), 20 μm (B), 4 μm (C, E), 2 μm (D, F, G, H, I, J).

图5 曼纳角毛藻的光学显微镜(LM) (A, I)、扫描电镜(SEM) (B-E, J)和透射电镜(TEM) (F-H)示意图。A: 链状群体; B: 链中壳面; C: 端壳面, 示唇形突(箭头); D: 链中壳面, 示重叠硅质翼(宽箭型), 硅质脊(箭头)及壳套基部凹槽(窄箭型); E-F: 链中壳面, 示中央环纹(F, 箭型); G: 壳套; H: 环带; I, J: 角毛结构。标尺 = 50 μm (A), 10 μm (B, D, I), 5 μm (F), 4 μm (C, E, J), 2 μm (G, H)。

Fig. 5 Morphology of Chaetoceros mannaii under light microscopy (LM) (A, I), scanning electron microscopy (SEM) (B-E, J) and transmission electron microscopy (TEM) (F-H). A, Chain in broad girdle view; B, Sibling intercalary valves showing aperture; C, Terminal valve view with external process of rimoportula (arrowhead); D, Intercalary cells with overlapping silica wings (broad arrow), silica ridges (arrowhead) and furrow above the basal ring of mantle (arrow); E, F, Internal view of an intercalary valves, showing central annulus (F, arrow); G, Mantle; H, Girdle bands; I and J: Setae structure. Scale bars, 50 μm (A), 10 μm (B, D, I), 5 μm (F), 4 μm (C, E, J), 2 μm (G, H).

图6 稀树角毛藻的光学显微镜(LM) (A-B)、扫描电镜(SEM) (C-D, I-J)和透射电镜(TEM) (E-H)示意图。A: 链状群体; B: 休眠孢子位于母细胞链中; C: 链中壳面, 示重叠硅质翼(箭型)及壳套基部凹槽(箭头); D: 释放的成熟休眠孢子; E: 链端壳面, 示中央唇形突(箭头)和硅质肋纹(箭型); F: 壳套; G: 链中壳面, 示中央环纹(箭型); H: 环带; I-J: 角毛结构。标尺 = 20 μm (A), 10 μm (D), 4 μm (C, D, E), 2 μm (F), 1 μm (G, H, I, J)。

Fig. 6 Morphology of Chaetoceros pauciramosus under light microscopy (LM) (A-B), scanning electron microscopy (SEM) (C-D, I-J) and transmission electron microscopy (TEM) (E-H). A, Chain in broad girdle view; B, Resting spores within the mother cells of a chain; C, Sibling intercalary valves showing overlapping silica ear-like structures (arrows) and furrow above the basal ring of mantle (arrowhead); D, Released resting spore; E, Terminal valve with short external tubes of rimoportulae (arrowhead) and silica rib (arrows); F, Mantle; G, Intercalary valve, showing central annulus (arrow); H, Girdle bands; I and J, Seta structure. Scale bars, 20 μm (A), 10 μm (D), 4 μm (C, D, E), 2 μm (F), 1 μm (G, H, I, J).

图7 基于核糖体大亚基部分序列的最大似然树。分支上的置信值分别显示贝叶斯分析和最大似然分析。

Fig. 7 Molecular phylogenetic tree inferred from LSU rDNA, with Chaetoceros diadema as outgroup. Supporting values on each nodule are from Bayesian and Maximum Likelihood analysis.

表1 洛氏角毛藻复合群中各个物种之间的遗传距离(括号内为碱基差异数)

Table 1 Genetic distance among allied taxa within the Chaetoceros lorenzianus complex (data in the brackets are numbers of different base pairs)

	平孢角毛藻 C. laevisporus	密特拉角毛藻 C. mitra	曼纳角毛藻 C. mannaii	并基角毛藻 C. decipiens	优美角毛藻 C. elegans
密特拉角毛藻 C. mitra	0.081 (52)
曼纳角毛藻 C. mannaii	0.105 (69)	0.047 (33)
并基角毛藻 C. decipiens	0.089 (58)	0.028 (20)	0.057 (38)
优美角毛藻 C. elegans	0.083 (53)	0.036 (25)	0.067 (44)	0.027 (17)
稀树角毛藻 C. pauciramosus	0.092 (60)	0.040 (26)	0.059 (39)	0.028 (18)	0.028 (21)

表2 洛氏角毛藻复合群相似物种之间的形态学比较

Table 2 Morphological comparison among allied taxa within Chaetoceros lorenzianus complex

特征 Character	并基角毛藻 C. decipiens	优美角毛藻 C. elegans	平孢角毛藻 C. laevisporus	曼纳角毛藻 C. mannaii	稀树角毛藻 C. pauciramosus	密特拉角毛藻 C. mitra	洛氏角毛藻 C. lorenzianus type material
角毛孔纹形状 Seta poroid shape	椭圆形 Oval	水滴状 Drop-shaped	椭圆形 Oval	椭圆形 Oval	细长形 Elongated	圆形或椭圆形 Round or oval	圆形 Oval
角毛孔纹大小 Seta poroid size (μm)	0.3-0.6 (0.4 ± 0.1)	0.3-1.6 (0.7 ± 0.3)	0.3-0.9 (0.6 ± 0.1)	0.8-1.5 (1.1 ± 0.1)	0.1-0.6 (0.3 ± 0.1)	0.1-0.3 (0.2 ± 0.1)	Nd Nd
角毛孔纹密度 Seta poroid number in 10 μm	14-25 (20.2 ± 3.6)	6-27 (18.7 ± 5.0)	11-17 (14.1 ± 1.9)	6-10 (7.9 ± 1.0)	18-44 (31.7 ± 5.4)	30-56 (39.8 ± 7.4)	5-9 (7.2 ± 1.7)
Brunel 型 Brunel group	I I	I I	I I	I I	I I	II II	I I
角毛基部并行融合Fusion of seta bases	有或无 Present/absent	无 Absent	无 Absent	无 Absent	短或无 Short/absent	无 Absent	有 Present
休眠孢子 Resting spore	未发现 Unknown	有二叉分支 Two branching processes	壳面平滑 Smooth	未发现 Unknown	有二叉分支 Two branching processes	有二叉分支 Two branching processes	有二叉分支? Two branching processes?
窗孔形状 Aperture shape	椭圆形 Oval	圆形或四边形 Rounded or quadrangular	椭圆形 Oval	六边形 Hexagonal	六边形或花生形 Hexagonal or peanut shaped	六边形或花生形Hexagonal or peanut shaped	圆形或六边形 Oval or hexagonal
角毛基部 Basal part of setae	无 Lacking	有且明显 Distinct	无 Lacking	短 Short	短 Short	无 Lacking	无 Lacking
壳面及壳套孔纹 Poroids on valve face and mantle	有 Yes	有 Yes	无 No	无 No	有 Yes	无 No	nd nd

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中国沿海洛氏角毛藻复合群的多样性组成及地理分布

Species diversity and geographical distribution of the Chaetoceros lorenzianus complex along the coast of China

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