生物多样性 ›› 2019, Vol. 27 ›› Issue (2): 149-158. DOI: 10.17520/biods.2018261
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:
Chen Zuoyi1,2,Xu Xiaojing1,Zhu Suying1,Zhai Mengyi1,Li Yang1,*()
Received:
2018-09-28
Accepted:
2019-01-27
Online:
2019-02-20
Published:
2019-04-16
Contact:
Li Yang
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。
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.
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.
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) |
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 |
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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