中国沿海洛氏角毛藻复合群的多样性组成及地理分布
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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。
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.
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.
引文
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)[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.]
Chen ZY,Lundholm N,Moestrup?,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.
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.]
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.]
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.]
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.]
Evensen DL,Hasle GR(1975)The morphology of some Chaetoceros(Bacillariophyceae)species as seen in the electron microscopes.Nova Hedwigia,53,152-174.
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.
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.]
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.]
Hall TA(1999)BioEdit:A user-friendly biological sequence alignment editor and analysis program for Window95/98/NT.Nucleic Acids Symposium Series,41,95-98.
Hasle GR,Syvertsen EE(1997)Marine diatoms.In:Identifying Marine Phytoplankton(ed.Tomas CR),pp.5-387.Academic Press,London.
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.
Jensen KG,Moestrup ?(1998)The genus Chaetoceros(Bacillariophyceae)in inner Danish coastal waters.Nordic Journal of Botany,18,88.
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.
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.
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.
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)[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.]
Lundholm N,Daugbjerg N,Moestrup ?(2002)Phylogeny of the Bacillariaceae with emphasis on the genus Pseudonitzschia(Bacillariophyceae)based on partial LSU rD NA.European Journal of Phycology,37,115?134.
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.
Nylander JAA(2004)MrModeltest v2.Program distributed by the author.Evolutionary Biology Center,Uppsala University,Uppsala,Sweden.
Okamura K(1911)Some littoral diatoms of Japan.Report Imperial Fisheries Institute Tokyo Japan.7,3-18.
Rines JEB,Hargraves PE(1988)The Chaetoceros Ehrenberg(Bacillariophyceae)flora of Narragansett Bay,Rhode Island,USA.Bible Phycology,79,5-196.
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.
Round FE,Crawford RM,Mann DG(1990)The Diatoms:Biology and Morphology of the Genera,p.747.Cambridge University Press,Cambridge.
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.]
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)[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.]
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.]
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.]
Chen ZY,Lundholm N,Moestrup?,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.
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.]
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.]
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.]
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.]
Evensen DL,Hasle GR(1975)The morphology of some Chaetoceros(Bacillariophyceae)species as seen in the electron microscopes.Nova Hedwigia,53,152-174.
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.
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.]
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.]
Hall TA(1999)BioEdit:A user-friendly biological sequence alignment editor and analysis program for Window95/98/NT.Nucleic Acids Symposium Series,41,95-98.
Hasle GR,Syvertsen EE(1997)Marine diatoms.In:Identifying Marine Phytoplankton(ed.Tomas CR),pp.5-387.Academic Press,London.
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.
Jensen KG,Moestrup ?(1998)The genus Chaetoceros(Bacillariophyceae)in inner Danish coastal waters.Nordic Journal of Botany,18,88.
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.
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.
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.
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)[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.]
Lundholm N,Daugbjerg N,Moestrup ?(2002)Phylogeny of the Bacillariaceae with emphasis on the genus Pseudonitzschia(Bacillariophyceae)based on partial LSU rD NA.European Journal of Phycology,37,115?134.
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.
Nylander JAA(2004)MrModeltest v2.Program distributed by the author.Evolutionary Biology Center,Uppsala University,Uppsala,Sweden.
Okamura K(1911)Some littoral diatoms of Japan.Report Imperial Fisheries Institute Tokyo Japan.7,3-18.
Rines JEB,Hargraves PE(1988)The Chaetoceros Ehrenberg(Bacillariophyceae)flora of Narragansett Bay,Rhode Island,USA.Bible Phycology,79,5-196.
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.
Round FE,Crawford RM,Mann DG(1990)The Diatoms:Biology and Morphology of the Genera,p.747.Cambridge University Press,Cambridge.
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.]
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)[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.]
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.]
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.]