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

• 研究报告 • 上一篇    下一篇

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. 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
  • 通讯作者: 李扬 E-mail:liyang@scnu.edu.cn
  • 基金项目:
    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 Zuoyi1, 2, Xu Xiaojing1, Zhu Suying1, Zhai Mengyi1, Li Yang1, *()   

  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
  • Contact: Li Yang E-mail:liyang@scnu.edu.cn

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

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

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。"

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)。"

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)。"

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)。"

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)。"

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)。"

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。"

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)"

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"

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

Absent

Absent

Absent
短或无
Short/absent

Absent

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

Lacking
有且明显
Distinct

Lacking

Short

Short

Lacking

Lacking
壳面及壳套孔纹
Poroids on valve
face and mantle

Yes

Yes

No

No

Yes

No
nd
nd
[1] Chen ZY, Li Y ( 2017) Preliminary study on some taxonomic puzzles of Chaetoceros decipiens Cleve. Acta Hydrobiologica Sinica, 41, 914-922. (in Chinese with English abstract)
doi: 10.7541/2017.114
[ 陈作艺, 李扬 ( 2017) 并基角毛藻若干分类学疑问的初步探讨. 水生生物学报, 41, 914-922.]
doi: 10.7541/2017.114
[2] Chen ZY, Lundholm N, Moestrup Ø, Kownacka J, Li Y ( 2018) Chaetoceros pauciramosus sp. nov. (Bacillariophyceae), a widely distributed brackish water species in the C. lorenzianus complex. Protist, 169, 615-631.
doi: 10.1016/j.protis.2018.06.007
[3] Chin TG, Chen JH, Huang KG ( 1965) Marine Planktonic Diatoms from China Sea, p. 115. Shanghai Science and Technology Press, Shanghai. (in Chinese)
[ 金德祥, 陈金环, 黄凯歌 ( 1965) 中国海洋浮游硅藻类, 见115页. 上海科学技术出版社, 上海.]
[4] Chin TG ( 1951) A list of Chinese diatoms from 1847 to 1946. Amoy Fishries Bulletin, 5, 145-230. (in Chinese with English abstract)
[ 金德祥 ( 1951) 中国硅藻目录. 厦门水产学报, 5, 145-230.]
[5] Chu SP, Kuo YC ( 1957) Studies on the genus Chaetoceros Ehrenberg from the fishing ground of the mackerel, Pneumatophorus japonicus (Houttuyn), off the Shantung coastal from Chefoo to Weihai. Part I. A systematic study. Oceanologia et Limnologia Sinica, 1, 27-87. (in Chinese with English abstract)
[ 朱树屏, 郭玉洁 ( 1957) 烟台、威海鲐鱼渔场及其附近海区角毛硅藻属的研究. I. 分类的研究. 海洋与湖沼, 1, 27-87.]
[6] Chu SP, Kuo YC ( 1958) Studies on the genus Chaetoceros Ehrenberg from the fishing ground of the mackerel, Pneumatophorus japonicus (Houttuyn), off the Shanung coast from Chefoo to Weihai. Part II. An ecological study. Oceanologia et Limnologia Sinica, 1, 167-179. (in Chinese with English abstract)
[ 朱树屏, 郭玉洁 ( 1958) 烟台、威海鲐鱼渔场及其附近海区角毛硅藻属的研究. II. 生态的研究. 海洋与湖沼, 1, 167-179.]
[7] Evensen DL, Hasle GR ( 1975) The morphology of some Chaetoceros (Bacillariophyceae) species as seen in the electron microscopes. Nova Hedwigia, 53, 152-174.
[8] Grunow A ( 1863) About some new and insufficiently known species and genera of diatoms. Negotiations of the Imperial Royal Zoological-Botanical Society in Vienna, 13, 137-162.
[9] Guo YJ ( 1963) The nature of Chaetoceros flora of the Yellow Sea. Oceanologia et Limnologia Sinica, 5, 322-332. (in Chinese with English abstract)
[ 郭玉洁 ( 1963) 黃海角毛藻属(Genus Chaetoceros Ehrenberg)区系的性质. 海洋与湖沼, 5, 322-332.]
[10] Guo YJ, Qian SB ( 2003) Flora Algarum Marinarum Sinicarum, Tomus V. Bacillariophyta, No.1 Centricae, pp. 345-346. Science Press, Beijing. (in Chinese)
[ 郭玉洁, 钱树本 ( 2003) 中国海藻志 (第5卷): 硅藻门 (第一册), 中心纲. 345-346页. 科学出版社, 北京.]
[11] Hall TA ( 1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for Window 95/98/NT. Nucleic Acids Symposium Series, 41, 95-98.
[12] Hasle GR, Syvertsen EE ( 1997) Marine diatoms. In: Identifying Marine Phytoplankton (ed. Tomas CR), pp. 5-387. Academic Press, London.
[13] Hernández-Becerril DU ( 1996) A morphological study of Chaetoceros species (Bacillariophyta) from the plankton of the Pacific Ocean of Mexico. Bulletin of the Natural History Museum (Botany series), 26, 1-73.
doi: 10.1128/JCM.01047-13
[14] Jensen KG, Moestrup ? ( 1998) The genus Chaetoceros (Bacillariophyceae) in inner Danish coastal waters. Nordic Journal of Botany, 18, 88.
doi: 10.1111/j.1756-1051.1998.tb01103.x
[15] Kooistra WHCF, Sarno D, Hernández-Becerril DU, Assmy P, Prisco CD, Montresor M ( 2010) Comparative molecular and morphological phylogenetic analyses of taxa in the Chaetocerataceae (Bacillariophyta). Phycologia, 49, 471-500.
doi: 10.2216/09-59.1
[16] Kownacka J, Edler L, Gromisz S, ?otocka M, Olenina I, Ostrowska M, Piwosz K ( 2013) Non-indigenous species Chaetoceros cf. lorenzianus Grunow 1863— A new, predominant component of autumn phytoplankton in the southern Baltic Sea. Estuarine Coastal & Shelf Science, 119, 101-111.
doi: 10.1016/j.ecss.2013.01.010
[17] Li Y, Boonprakob A, Gaonkar CC, Kooistra WHCF, Lange CB, Hernández-Becerril DU, Chen ZY, Moestrup ?, Lundholm N ( 2017) Diversity in the globally distributed diatom genus Chaetoceros (Bacillariophyceae): Three new species from warm-temperate waters. PLoS ONE, 12, 1-38.
[18] Lin GM, Yang QL ( 2007) Species diversity and the distribution of micro-phytoplankton in the Taiwan Strait. Biodiversity Science, 15, 31-45. (in Chinese with English abstract)
doi: 10.3321/j.issn:1005-0094.2007.01.003
[ 林更铭, 杨清良 ( 2007) 台湾海峡小型浮游植物的物种多样性和分布特征. 生物多样性, 15, 31-45.]
doi: 10.3321/j.issn:1005-0094.2007.01.003
[19] Lundholm N, Daugbjerg N, Moestrup ? ( 2002) Phylogeny of the Bacillariaceae with emphasis on the genus Pseudo- nitzschia (Bacillariophyceae) based on partial LSU rDNA. European Journal of Phycology, 37, 115-134.
doi: 10.1017/S096702620100347X
[20] Miller MA, Pfeiffer W, Schwartz T ( 2010) Creating the CIPRES science gateway for inference of large phylogenetic trees. 2010 Gateway Computing Environments Workshop (GCE), New Orleans, LA, pp.1-8.
doi: 10.1109/GCE.2010.5676129
[21] Nylander JAA ( 2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Center, Uppsala University, Uppsala, Sweden.
[22] Okamura K ( 1911) Some littoral diatoms of Japan. Report Imperial Fisheries Institute Tokyo Japan. 7, 3-18.
[23] Rines JEB, Hargraves PE ( 1988) The Chaetoceros Ehrenberg (Bacillariophyceae) flora of Narragansett Bay, Rhode Island, USA. Bible Phycology, 79, 5-196.
[24] Ronquist F, Teslenko M, Mark P, Ayres DL, Darling A ( 2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539-542.
doi: 10.1093/sysbio/sys029 pmid: 3329765
[25] Round FE, Crawford RM, Mann DG ( 1990) The Diatoms: Biology and Morphology of the Genera, p. 747. Cambridge University Press, Cambridge.
[26] Wang Y, Nie R, Li Y, Lü SH ( 2010) Species diversity and geographical distribution of Chaetoceros in Guangdong coast waters. Advance in Marine Science, 28, 342-352. (in Chinese with English abstract)
[ 王艳, 聂瑞, 李扬, 吕颂辉 ( 2010) 广东沿海角毛藻(Chaetoceros)的种类多样性及其地理分布. 海洋科学进展, 28, 342-352.]
[27] Xue B, Sun J, Li TT ( 2016) Phytoplankton community structure of northern South China Sea in summer of 2014. Haiyang Xuebao, 8, 54-65. (in Chinese with English abstract)
doi: 10.3969/j.issn.0253-4193.2016.04.005
[ 薛冰, 孙军, 李婷婷 ( 2016) 2014年夏季南海北部浮游植物群落结构.海洋学报, 8, 54-65.]
doi: 10.3969/j.issn.0253-4193.2016.04.005
[28] Yang Y, Sun J, Guan XY, Zhai WD, Guo SJ ( 2016) Seasonal variation of net-phytoplankton community in Bohai Sea. Marine Science Bulletin, 35, 121-131. (in Chinese with English abstract)
[ 杨阳, 孙军, 关翔宇, 翟惟东, 郭术津 ( 2016) 渤海网采浮游植物群集的季节变化. 海洋通报, 35, 121-131.]
[29] Zhai MY, Zhu SY, Chen ZY, Li Y ( 2017) Preliminary study on the species diversity of Chaetoceros lorenzianus complex from Guangdong coastal waters. Acta Hydrobiologica Sinica, 41, 1282-1290. (in Chinese with English abstract)
[ 翟梦怡, 朱素英, 陈作艺, 李扬 ( 2017) 广东沿海洛氏角毛藻复合群物种多样性的探究. 水生生物学报, 41, 1282-1290.]
[1] 邹安龙, 马素辉, 倪晓凤, 蔡琼, 李修平, 吉成均. 模拟氮沉降对北京东灵山辽东栎群落林下植物物种多样性的影响[J]. 生物多样性, 2019, 27(6): 607-618.
[2] 胡宜峰, 余文华, 岳阳, 黄正澜懿, 李玉春, 吴毅. 海南岛翼手目物种多样性现状与分布预测[J]. 生物多样性, 2019, 27(4): 400-408.
[3] 颜文博,吉晟男,帅凌鹰,赵雷刚,朱大鹏,曾治高. 秦岭南坡陕西洋县辖区哺乳动物物种多样性的空间分布格局[J]. 生物多样性, 2019, 27(2): 177-185.
[4] 邹东廷, 王庆刚, 罗奥, 王志恒. 中国蔷薇科植物多样性格局及其资源植物保护现状[J]. 植物生态学报, 2019, 43(1): 1-15.
[5] 王波, 黄勇, 李家堂, 戴强, 王跃招, 杨道德. 西南喀斯特地貌区两栖动物丰富度分布格局与环境因子的关系[J]. 生物多样性, 2018, 26(9): 941-950.
[6] 孙德鑫, 刘向, 周淑荣. 停止人为去除植物功能群后的高寒草甸多样性恢复过程与群落构建[J]. 生物多样性, 2018, 26(7): 655-666.
[7] 张宇, 冯刚. 内蒙古昆虫物种多样性分布格局及其机制[J]. 生物多样性, 2018, 26(7): 701-706.
[8] 罗俊杰, 王莹, 商辉, 周喜乐, 韦宏金, 黄素楠, 顾钰峰, 金冬梅, 戴锡玲, 严岳鸿. 基于孢子形态和分子证据探讨鳞盖蕨属(碗蕨科)系统分类[J]. 植物学报, 2018, 53(6): 782-792.
[9] 陶夏秋, 崔绍朋, 蒋志刚, 初红军, 李娜, 杨道德, 李春旺. 新疆阿勒泰地区爬行动物区系及多样性海拔分布格局[J]. 生物多样性, 2018, 26(6): 578-589.
[10] 田成, 李俊清, 杨旭煜, 余鳞, 袁丹, 黎运喜. 利用红外相机技术对四川王朗国家级自然保护区野生动物物种多样性的初步调查[J]. 生物多样性, 2018, 26(6): 620-626.
[11] 杨倩, 王娓, 曾辉. 氮添加对内蒙古退化草地植物群落多样性和生物量的影响[J]. 植物生态学报, 2018, 42(4): 430-441.
[12] 温韩东, 林露湘, 杨洁, 胡跃华, 曹敏, 刘玉洪, 鲁志云, 谢有能. 云南哀牢山中山湿性常绿阔叶林20 hm2动态样地的物种组成与群落结构[J]. 植物生态学报, 2018, 42(4): 419-429.
[13] 刘海跃, 李欣玫, 张琳琳, 王姣姣, 贺学礼. 西北荒漠带花棒根际丛枝菌根真菌生态地理分布[J]. 植物生态学报, 2018, 42(2): 252-260.
[14] 胡一鸣, 梁健超, 金崑, 丁志锋, 周智鑫, 胡慧建, 蒋志刚. 喜马拉雅山哺乳动物物种多样性垂直分布格局[J]. 生物多样性, 2018, 26(2): 191-201.
[15] 商天其, 叶诺楠, 高海卿, 高洪娣, 伊力塔. 基于多元回归树的公益林群落结构解析[J]. 植物学报, 2018, 53(2): 238-249.
Viewed
Full text


Abstract

Cited

  Shared   
  Discussed