生物多样性 ›› 2015, Vol. 23 ›› Issue (3): 383-397. DOI: 10.17520/biods.2014252
宋会银1,2, 张琪1, 胡愈炘1,2, 刘国祥1,,A;*, 胡征宇1
收稿日期:
2014-12-03
接受日期:
2015-03-19
出版日期:
2015-06-08
发布日期:
2015-06-12
通讯作者:
刘国祥
基金资助:
Huiyin Song1,2, Qi Zhang1, Yuxin Hu1,2, Guoxiang Liu1,*, Zhengyu Hu1
Received:
2014-12-03
Accepted:
2015-03-19
Online:
2015-06-08
Published:
2015-06-12
Contact:
Liu Guoxiang
摘要:
球状绿藻主要指绿藻门中多为单细胞, 形状为球形、近球形, 或由球形衍生出来的其他形状的藻类。球状绿藻分布广泛, 遍布全球, 生活于淡水、海水和亚气生等生境中, 其相似的简单形态下隐藏着复杂的物种多样性。球状绿藻分类学上主要位于绿藻门的两纲四目及其一些独立支系的类群。球状绿藻分类学正由传统的基于形态特征向基于分子信息的复合分类方法转变。球状绿藻隐性的物种多样性涉及约40属, 其中15属是依据新标本材料而建立的新属, 12个属是依据新证据建立的新组合。本文重点介绍了1998年以后在共球藻纲和绿藻纲中新发现和命名的单细胞球状绿藻, 介绍了它们的形态特征、分类学迁移及理由, 特别对常见的小球藻属和栅藻属的分类学概念变迁作了详细介绍, 对一些尚没有中文名的拉丁学名给出了中文命名。另外, 本文讨论了基于DNA的分类学方法在球状绿藻分类学中的应用, 目前系统发育位置的不同已经成为球状绿藻分类的主要依据。目前球状绿藻分类面临的问题是大多分类位置未用分子系统发育方法的验证。 未来, 球状绿藻分类学家应用系统发育研究结合形态学研究探索单细胞球状绿藻的生物多样性。
宋会银, 张琪, 胡愈炘, 刘国祥, 胡征宇 (2015) 球状绿藻的隐性生物多样性及其分类学进展. 生物多样性, 23, 383-397. DOI: 10.17520/biods.2014252.
Huiyin Song, Qi Zhang, Yuxin Hu, Guoxiang Liu, Zhengyu Hu (2015) Cryptic biodiversity of coccoid green algae and progress in the phylogenic studies. Biodiversity Science, 23, 383-397. DOI: 10.17520/biods.2014252.
绿藻门 Chlorophyta |
---|
共球藻纲 Trebouxiophyceae |
渡边球藻分支 The Watanabea clade |
椭圆球藻属 Chloroidium Nadson 沙漠球藻属 Desertella Fucikova et al. 异小球藻属 Heterochlorella Neustupa et al. 胶树小球藻属 Heveochlorella Zhang et al. 卡莉娜球藻属 Kalinella Neustupa Elias et al. 假椭圆球藻属 Parachloroidium Neustupa et Skaloud 鲜绿球藻属 Viridiella Albertano et al. 渡边球藻属 Watanabea Hanagata et al. |
共球藻目 Trebouxiales |
星绿藻属 Asterochloris Tschermak-Woess 共球藻属 Trebouxia Puymaly |
小球藻目 Chlorellales 小球藻分支 Chlorella-clade |
集星藻属 Actinastrum Lagerheim 小球藻属 Chlorella Beyerinck 对囊藻属 Didymogenes Schmidle 黑氏球藻属 Hegewaldia Proschold 亨氏球藻属 Hindakia Bock et al. 海尼球藻属 Heynigia Bock et al. 迈耶球藻属 Meyerella Fawley et al. 微芒藻属 Micractinium Fresinius 类小球藻分支 Parachlorella-clade 胶网藻属 Dictyosphaerium Nageli 霉球藻属 Mucidosphaerium Bock et al. 类小球藻属 Parachlorella Krienitz et al. 浮小球藻属 Planktochlorella Skaloud et Nemcova |
共球藻纲内形成独立支系的属 Incertae sedis in Trebouxiophyceae |
绿微藻属 Chloroparva Somogyi et al. 绿梨藻属 Chloropyrula Gaysina et al. 胶球藻属 Coccomyxa Schmidle 独绿藻属 Eremochloris Fucikova et al.. 微小球藻属 Leptochlorella Neustupa et al. 干球藻属 Xerochlorella Fucikova et al. 丛林球藻属 Xylochloris Neustupa et al. |
绿藻纲 Chlorophyceae |
环藻目 Sphaeropleales |
栅藻属 Scenedesmus Meyen 链带藻属 Desmodesmus (Chodat) An et al. 尖带藻属 Acutodesmus (Hegewald) Tsarenko 乔带藻属 Chodatodesmus Hegewald et al. 疣带藻属 Verrucodesmus Hegewald 木带藻属 Hylodesmus Elias et al. 扇带藻属 Pectinodesmus Hegewald et al. 考马斯藻属 Comasiella Hegewald et al. |
片球藻属 Bracteacoccus Tereg 色绿藻属 Chromochloris Kol & Chodat |
麦可藻属 Mychonastes Simpson & Van Valkenburg 衣藻目 Chlamydomonadales |
拟博罗德藻属 Borodinellopsis Dykstra 绿球藻属 Chlorococcum Meneghini |
绿藻纲中形成独立支系的属 Incertae sedis in Chlorophyceae 耶氏球藻属 Jenufa Nemcova et al. |
表1 文中介绍的球状绿藻的分类名录
Table 1 Checklist of the coccoid green algae in the present study
绿藻门 Chlorophyta |
---|
共球藻纲 Trebouxiophyceae |
渡边球藻分支 The Watanabea clade |
椭圆球藻属 Chloroidium Nadson 沙漠球藻属 Desertella Fucikova et al. 异小球藻属 Heterochlorella Neustupa et al. 胶树小球藻属 Heveochlorella Zhang et al. 卡莉娜球藻属 Kalinella Neustupa Elias et al. 假椭圆球藻属 Parachloroidium Neustupa et Skaloud 鲜绿球藻属 Viridiella Albertano et al. 渡边球藻属 Watanabea Hanagata et al. |
共球藻目 Trebouxiales |
星绿藻属 Asterochloris Tschermak-Woess 共球藻属 Trebouxia Puymaly |
小球藻目 Chlorellales 小球藻分支 Chlorella-clade |
集星藻属 Actinastrum Lagerheim 小球藻属 Chlorella Beyerinck 对囊藻属 Didymogenes Schmidle 黑氏球藻属 Hegewaldia Proschold 亨氏球藻属 Hindakia Bock et al. 海尼球藻属 Heynigia Bock et al. 迈耶球藻属 Meyerella Fawley et al. 微芒藻属 Micractinium Fresinius 类小球藻分支 Parachlorella-clade 胶网藻属 Dictyosphaerium Nageli 霉球藻属 Mucidosphaerium Bock et al. 类小球藻属 Parachlorella Krienitz et al. 浮小球藻属 Planktochlorella Skaloud et Nemcova |
共球藻纲内形成独立支系的属 Incertae sedis in Trebouxiophyceae |
绿微藻属 Chloroparva Somogyi et al. 绿梨藻属 Chloropyrula Gaysina et al. 胶球藻属 Coccomyxa Schmidle 独绿藻属 Eremochloris Fucikova et al.. 微小球藻属 Leptochlorella Neustupa et al. 干球藻属 Xerochlorella Fucikova et al. 丛林球藻属 Xylochloris Neustupa et al. |
绿藻纲 Chlorophyceae |
环藻目 Sphaeropleales |
栅藻属 Scenedesmus Meyen 链带藻属 Desmodesmus (Chodat) An et al. 尖带藻属 Acutodesmus (Hegewald) Tsarenko 乔带藻属 Chodatodesmus Hegewald et al. 疣带藻属 Verrucodesmus Hegewald 木带藻属 Hylodesmus Elias et al. 扇带藻属 Pectinodesmus Hegewald et al. 考马斯藻属 Comasiella Hegewald et al. |
片球藻属 Bracteacoccus Tereg 色绿藻属 Chromochloris Kol & Chodat |
麦可藻属 Mychonastes Simpson & Van Valkenburg 衣藻目 Chlamydomonadales |
拟博罗德藻属 Borodinellopsis Dykstra 绿球藻属 Chlorococcum Meneghini |
绿藻纲中形成独立支系的属 Incertae sedis in Chlorophyceae 耶氏球藻属 Jenufa Nemcova et al. |
绿藻门Chlorophyta | |
---|---|
绿藻纲 Chlorophyceae | 共球藻纲 Trebouxiophyceae |
衣藻目/团藻目 Chlamydomonadales* | 共球藻目 Trebouxiales* |
环藻目 Sphaeropleales* | 渡边球藻分支 Watanabea clade* |
鞘藻目 Oedogoniales | 溪菜目 Prasiolales |
楯毛藻目 Chaetopeltidales | 小球藻目Chlorellales* |
胶毛藻目 Chaetophorales | 绿囊球藻/葡萄藻分支Choricystis/Botryococcus clade* |
分类位置未确定的类群Incertae sedis* | 分类位置未确定的类群Incertae sedis* |
附录1 球藻绿藻的分类学位置
Appendix 1 Phylogeny of the coccoid green algae
绿藻门Chlorophyta | |
---|---|
绿藻纲 Chlorophyceae | 共球藻纲 Trebouxiophyceae |
衣藻目/团藻目 Chlamydomonadales* | 共球藻目 Trebouxiales* |
环藻目 Sphaeropleales* | 渡边球藻分支 Watanabea clade* |
鞘藻目 Oedogoniales | 溪菜目 Prasiolales |
楯毛藻目 Chaetopeltidales | 小球藻目Chlorellales* |
胶毛藻目 Chaetophorales | 绿囊球藻/葡萄藻分支Choricystis/Botryococcus clade* |
分类位置未确定的类群Incertae sedis* | 分类位置未确定的类群Incertae sedis* |
属名 | 分类依据 | 主要分类方法 | ||
---|---|---|---|---|
Chloroidium Nadson, 1906 (Darienko等(2010)重新提出) | 新证据(从小球藻属独立) | 基于形态、生殖、生态生理和SSU、ITS系统发育研究 | ||
Parachloroidium Neustupa, 2013 | 新标本 | 基于SSU、ITS和rbcL序列的系统发育研究 | ||
Kalinella Neustupa et al., 2009 | 新标本 | 基于SSU和rbcL序列系统发育研究结合形态观察 | ||
Heterochlorella Neustupa et al., 2009 | 新证据(从小球藻属独立) | 基于SSU系统发育研究 | ||
Heveochlorella Zhang et al., 2008 | 新证据(从小球藻属独立) | 基于SSU和16S rDNA序列系统发育研究结合形态观察 | ||
Watanabea Hanagata et al., 1998 | 新标本 | 基于形态特征 | ||
Desertella | 新标本 | 基于SSU 和rbcL序列系统发育研究 | ||
Meyerella Fawley et al., 2005 | 新标本 | 基于SSU、ITS和rbcL序列的系统发育研究 | ||
Hegewaldia Proschold et al., 2010 | 新证据(从微芒藻属独立) | 基于SSU和ITS rDNA的系统发育研究 | ||
Hindakia Bock et al., 2010 | 新证据(从胶网藻属独立) | 基于SSU 和ITS rDNA的系统发育研究 | ||
Heynigia Bock et al., 2010 | 新标本 | 基于SSU 和ITS rDNA的系统发育研究 | ||
Parachlorella Krieniz et al., 2004 | 新证据(从小球藻属独立) | 基于SSU 和ITS rDNA系统发育研究二级结构研究 | ||
Mucidosphaerium Bock et al., 2011 | 新证据(从胶网藻属独立) | 基于SSU系统发育、ITS rDNA二级结构和形态分析 | ||
Planktochlorella Skaloud, 2014 | 新标本 | 基于SSU 系统发育和ITS2二级结构研究 | ||
Xylochloris Neustupa et al., 2011 | 新标本 | 基于SSU系统发育研究 | ||
Chloroparva Somogyi et al., 2011 | 新标本 | 基于SSU系统发育研究 | ||
Chloropyrula Gaysina et al., 2013 | 新标本 | 基于SSU系统发育研究 | ||
Leptochlorella Neustupa et al., 2013 | 新标本 | 基于SSU 和rbcL序列系统发育研究 | ||
Eremochloris | 新标本 | 基于SSU 和rbcL序列系统发育研究 | ||
Xerochlorella | 新标本 | 基于SSU 和rbcL序列系统发育研究 | ||
Desmodesmus (Chodat) An et al., 1999 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Pectinodesmus Hegewald et al., 2010 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Comasiella Hegewald et al., 2010 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Chodatodesmus Hegewald et al., 2013 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Verrucodesmus Hegewald, 2013 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Hylodesmus Elias et al., 2009 | 新标本 | 基于SSU系统发育研究和 ITS2序列 | ||
Jenufa Nemcova et al., 2011 | 新标本 | 基于SSU系统发育研究 |
附录2 1998-2014年提出的球藻绿藻属及其分类依据
Appendix 2 The coccoid green algae genus proposed between 1998 and 2014 and their taxonic basis
属名 | 分类依据 | 主要分类方法 | ||
---|---|---|---|---|
Chloroidium Nadson, 1906 (Darienko等(2010)重新提出) | 新证据(从小球藻属独立) | 基于形态、生殖、生态生理和SSU、ITS系统发育研究 | ||
Parachloroidium Neustupa, 2013 | 新标本 | 基于SSU、ITS和rbcL序列的系统发育研究 | ||
Kalinella Neustupa et al., 2009 | 新标本 | 基于SSU和rbcL序列系统发育研究结合形态观察 | ||
Heterochlorella Neustupa et al., 2009 | 新证据(从小球藻属独立) | 基于SSU系统发育研究 | ||
Heveochlorella Zhang et al., 2008 | 新证据(从小球藻属独立) | 基于SSU和16S rDNA序列系统发育研究结合形态观察 | ||
Watanabea Hanagata et al., 1998 | 新标本 | 基于形态特征 | ||
Desertella | 新标本 | 基于SSU 和rbcL序列系统发育研究 | ||
Meyerella Fawley et al., 2005 | 新标本 | 基于SSU、ITS和rbcL序列的系统发育研究 | ||
Hegewaldia Proschold et al., 2010 | 新证据(从微芒藻属独立) | 基于SSU和ITS rDNA的系统发育研究 | ||
Hindakia Bock et al., 2010 | 新证据(从胶网藻属独立) | 基于SSU 和ITS rDNA的系统发育研究 | ||
Heynigia Bock et al., 2010 | 新标本 | 基于SSU 和ITS rDNA的系统发育研究 | ||
Parachlorella Krieniz et al., 2004 | 新证据(从小球藻属独立) | 基于SSU 和ITS rDNA系统发育研究二级结构研究 | ||
Mucidosphaerium Bock et al., 2011 | 新证据(从胶网藻属独立) | 基于SSU系统发育、ITS rDNA二级结构和形态分析 | ||
Planktochlorella Skaloud, 2014 | 新标本 | 基于SSU 系统发育和ITS2二级结构研究 | ||
Xylochloris Neustupa et al., 2011 | 新标本 | 基于SSU系统发育研究 | ||
Chloroparva Somogyi et al., 2011 | 新标本 | 基于SSU系统发育研究 | ||
Chloropyrula Gaysina et al., 2013 | 新标本 | 基于SSU系统发育研究 | ||
Leptochlorella Neustupa et al., 2013 | 新标本 | 基于SSU 和rbcL序列系统发育研究 | ||
Eremochloris | 新标本 | 基于SSU 和rbcL序列系统发育研究 | ||
Xerochlorella | 新标本 | 基于SSU 和rbcL序列系统发育研究 | ||
Desmodesmus (Chodat) An et al., 1999 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Pectinodesmus Hegewald et al., 2010 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Comasiella Hegewald et al., 2010 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Chodatodesmus Hegewald et al., 2013 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Verrucodesmus Hegewald, 2013 | 新证据(从栅藻属独立) | 基于ITS2系统发育研究 | ||
Hylodesmus Elias et al., 2009 | 新标本 | 基于SSU系统发育研究和 ITS2序列 | ||
Jenufa Nemcova et al., 2011 | 新标本 | 基于SSU系统发育研究 |
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