部分雀形目鸟类的CoⅠ和Cytb基因序列测定及其系统发育研究
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摘要
雀形目(Passeriformes);是现存鸟类中种类最多、适应辐射最广的类群,其系统发育关系一直存有争议。已有学者采用采用线粒体DNA等分子标记对部分雀形目鸟类的分子系统发育关系进行了一些探讨,但种类有限。本文通过PCR扩增、直接测序或克隆测序,对来自四川地区的部分雀形目鸟类的线粒体DNA细胞色素b基因全序列(1143 bp)和CoⅠ基因部分序列(1176 bp)的组成特点及其系统发育关系进行了比较分析,尤其针对分类地位上目前还存在争议的鸫亚科、画眉亚科鸟类,分别以树麻雀(Passer montanus)、荒漠伯劳(Lanius isabellinus)和灰树鹊(Dendrocitta formosae)为外群构建它们的系统发生树,深入探讨了这两个亚科鸟类的系统发育关系及其分类地位。主要结果如下：
1)获得了69种雀形目鸟类的Cytb基因全序列和CoⅠ基因部分序列,联合数据集序列长度为2319 bp,共有变异位点1000个,占总位点数的43.1%；简约信息位点923个,占变异位点数的92.3%；T、C、A、G碱基的平均含量分别为24.8%、32.2%、27.6%、15.4%。
2)Sibley & Ahlquist分类系统中雀小目(Corvida)的莺总科(Sylvioidea)、鹟总科(Muscicapoidea)和雀总科(Pasesoridea)三个类群,以及鸦小目(Passerida)的鸦总科(Corvoidea),在本文中得到进一步验证,但莺总科为非单系类群。山雀属(Parus)为一单独类群；长尾山雀类与莺亚科鸟类的关系远比与山雀属鸟类的关系近,不是山雀科(Paridae)的一个属。攀雀科(Remizidae)应归属于莺总科。燕雀类与鹀类的分歧达到科级水平,分别归属为燕雀科和鹀科。
3)鸫亚科鸟类为非单系,被分为2个支系,一个支系包括鸫属(Turdus)和地鸫属(Zoothera)。另一个支系包括红尾鸲属(Phoeicurus)、鸲属(Tarsiger)、燕尾属(Enicurus)和啸鸫属(Myiophonus)。宝兴歌鸫(Turdus mupinensis)独立于鸫属而与虎斑地鸫(Zoothera dauma)形成单独一个分支,宝兴歌鸫的分类地位独特。
4)画眉亚科鸟类被分为2大支系,非单系。传统认为属于画眉科的鵙鹛属(Pteruthius)、鳞胸鹪鹛(Pnoepyga albiventer)和褐胁雀鹛(Alcippe dubia)与画眉亚科其他物种之间亲缘关系很远,而暗绿绣眼鸟(Zosterops japonicus)置于画眉亚科中,其分类地位需要重新认识。并证实噪鹛属(Garrulax)和雀鹛属(Alcippe)为多系类群。鸦雀属(Paradoxornis)属于画眉亚科,穗鹛(Stachyris)和钩嘴鹛(Pomatorhinus)具有较近的亲缘关系,联合数据集未能较好地反映希鹛属物种在形态上的差异。
5)研究表明CoⅠ基因能够为重建鸟类系统发育关系提供足够的系统发育信息。基于与联合数据集的比较分析,COⅠ基因适合用于分析科内属间系统发育关系,当然它也能作为鉴别物种的分子标记,但不够稳定、准确,因此,本文建议应联合其他分子标记组合分析。
The birds of Passeriformes are the group which are the largest number and the broadest adaptive radiation of existent aves, their molecular phylogeny have been focused on for a long time. Former molecular phylogeny of the limited passerines had been studied by mitochondrial DNA. In this paper, by PCR amplification, direct sequencing or cloning and sequencing method, partion cytochrome oxidase subunitⅠgene (CoⅠ)and complete cytochrome b gene (Cytb) sequences of our samples in Sichuan were analyzed. We analyzed the sequences composition and characteristic of the combined sequences, and builted phylogenetic trees. Furthermore, Passer montanus, Lanius isabellinus and Dendrocitta formosae were respectively selected as outgroup, we deeply analysed their phylogenetic relationship to defecate the mistiness questions about tradition classify in the Turdinae and Timaliinae, and discussed the station of their species. The results showed as follow:
1)The combined sequences of the Cytb and CoⅠgenes of 69 species from songbird (Aves:Passeriformes) resulted in an alignment of 2319 nucleotides. The variable sites were 1000,43.1% of all sites. The parsimony informative sites were 923,92.3% of the variable sites. The average composition of four bases T, C, A and G were 24.8%,32.2%,27.6%, 15.4%, respectively.
2) This verified four superfamilies, i.e. Sylvioidea, Muscicapoidea and Pasesoridea within Passerida and Corvoidea within Passerida from Sibley-Ahlquist taxonomy based on DNA hybridization studies, but The Sylvioidea was nonmonophyly among them. The Paridae was a individual group, the Remizidae was placed in the superfamily Sylvioidea, This study also found that Aegithalos concinnus was distantly related to the genera Parus in the Paridae birds, but closely related to the Sylviinae.Fringillids and Emberizids reached a familial level and should be listed into the families Fringillidae and Emberizidae, respectively.
3)The Turdinae was nonmonophyly. In the phylogenetic trees, the species examined in the study were clustered into two clades. One clade included the genera Turdus and Zoothera. The other clade included the genera Phoenicurus, Enicurus, Tarsiger and Myiophonus. Turdus mupinensis was out of the genus Turdus, forming a single clade with Zoothera dauma, so its Taxonomic status was unusual.
4) The Timaliinae was nonmonophyly too. The results indicated that it consisted of 2 lineages. And several species traditionally placed among the Timaliinae, two species from the genus Pteruthius, Pnoepyga albiventer and Alcippe dubia, were not related to the Timaliinae. Furthermore, the phylogenetic hypotheses inferred from molecular data suggested that the Timaliinae should include Zosterops japonicas. All topologies of the genera in this study were stable, and confirmed that the genera Garrulax and Alcippe were polyphyletic groups. The Paradoxornis was a genus of the Timaliinae. The genera Stachyris and Pomatorhinus were closely related. The combined data could not indicate the morphological difference of species from the genus Minla.
5) Studies showed that the CoⅠgene was(suitable to identify the phylogenetic relationship of avian genus unit, and it can be used to identify avian species, but it was less stable and accurate than the combined sequences,so we suggested it should be analysed with other molecular makers.
1)获得了69种雀形目鸟类的Cytb基因全序列和CoⅠ基因部分序列,联合数据集序列长度为2319 bp,共有变异位点1000个,占总位点数的43.1%；简约信息位点923个,占变异位点数的92.3%；T、C、A、G碱基的平均含量分别为24.8%、32.2%、27.6%、15.4%。
2)Sibley & Ahlquist分类系统中雀小目(Corvida)的莺总科(Sylvioidea)、鹟总科(Muscicapoidea)和雀总科(Pasesoridea)三个类群,以及鸦小目(Passerida)的鸦总科(Corvoidea),在本文中得到进一步验证,但莺总科为非单系类群。山雀属(Parus)为一单独类群；长尾山雀类与莺亚科鸟类的关系远比与山雀属鸟类的关系近,不是山雀科(Paridae)的一个属。攀雀科(Remizidae)应归属于莺总科。燕雀类与鹀类的分歧达到科级水平,分别归属为燕雀科和鹀科。
3)鸫亚科鸟类为非单系,被分为2个支系,一个支系包括鸫属(Turdus)和地鸫属(Zoothera)。另一个支系包括红尾鸲属(Phoeicurus)、鸲属(Tarsiger)、燕尾属(Enicurus)和啸鸫属(Myiophonus)。宝兴歌鸫(Turdus mupinensis)独立于鸫属而与虎斑地鸫(Zoothera dauma)形成单独一个分支,宝兴歌鸫的分类地位独特。
4)画眉亚科鸟类被分为2大支系,非单系。传统认为属于画眉科的鵙鹛属(Pteruthius)、鳞胸鹪鹛(Pnoepyga albiventer)和褐胁雀鹛(Alcippe dubia)与画眉亚科其他物种之间亲缘关系很远,而暗绿绣眼鸟(Zosterops japonicus)置于画眉亚科中,其分类地位需要重新认识。并证实噪鹛属(Garrulax)和雀鹛属(Alcippe)为多系类群。鸦雀属(Paradoxornis)属于画眉亚科,穗鹛(Stachyris)和钩嘴鹛(Pomatorhinus)具有较近的亲缘关系,联合数据集未能较好地反映希鹛属物种在形态上的差异。
5)研究表明CoⅠ基因能够为重建鸟类系统发育关系提供足够的系统发育信息。基于与联合数据集的比较分析,COⅠ基因适合用于分析科内属间系统发育关系,当然它也能作为鉴别物种的分子标记,但不够稳定、准确,因此,本文建议应联合其他分子标记组合分析。
The birds of Passeriformes are the group which are the largest number and the broadest adaptive radiation of existent aves, their molecular phylogeny have been focused on for a long time. Former molecular phylogeny of the limited passerines had been studied by mitochondrial DNA. In this paper, by PCR amplification, direct sequencing or cloning and sequencing method, partion cytochrome oxidase subunitⅠgene (CoⅠ)and complete cytochrome b gene (Cytb) sequences of our samples in Sichuan were analyzed. We analyzed the sequences composition and characteristic of the combined sequences, and builted phylogenetic trees. Furthermore, Passer montanus, Lanius isabellinus and Dendrocitta formosae were respectively selected as outgroup, we deeply analysed their phylogenetic relationship to defecate the mistiness questions about tradition classify in the Turdinae and Timaliinae, and discussed the station of their species. The results showed as follow:
1)The combined sequences of the Cytb and CoⅠgenes of 69 species from songbird (Aves:Passeriformes) resulted in an alignment of 2319 nucleotides. The variable sites were 1000,43.1% of all sites. The parsimony informative sites were 923,92.3% of the variable sites. The average composition of four bases T, C, A and G were 24.8%,32.2%,27.6%, 15.4%, respectively.
2) This verified four superfamilies, i.e. Sylvioidea, Muscicapoidea and Pasesoridea within Passerida and Corvoidea within Passerida from Sibley-Ahlquist taxonomy based on DNA hybridization studies, but The Sylvioidea was nonmonophyly among them. The Paridae was a individual group, the Remizidae was placed in the superfamily Sylvioidea, This study also found that Aegithalos concinnus was distantly related to the genera Parus in the Paridae birds, but closely related to the Sylviinae.Fringillids and Emberizids reached a familial level and should be listed into the families Fringillidae and Emberizidae, respectively.
3)The Turdinae was nonmonophyly. In the phylogenetic trees, the species examined in the study were clustered into two clades. One clade included the genera Turdus and Zoothera. The other clade included the genera Phoenicurus, Enicurus, Tarsiger and Myiophonus. Turdus mupinensis was out of the genus Turdus, forming a single clade with Zoothera dauma, so its Taxonomic status was unusual.
4) The Timaliinae was nonmonophyly too. The results indicated that it consisted of 2 lineages. And several species traditionally placed among the Timaliinae, two species from the genus Pteruthius, Pnoepyga albiventer and Alcippe dubia, were not related to the Timaliinae. Furthermore, the phylogenetic hypotheses inferred from molecular data suggested that the Timaliinae should include Zosterops japonicas. All topologies of the genera in this study were stable, and confirmed that the genera Garrulax and Alcippe were polyphyletic groups. The Paradoxornis was a genus of the Timaliinae. The genera Stachyris and Pomatorhinus were closely related. The combined data could not indicate the morphological difference of species from the genus Minla.
5) Studies showed that the CoⅠgene was(suitable to identify the phylogenetic relationship of avian genus unit, and it can be used to identify avian species, but it was less stable and accurate than the combined sequences,so we suggested it should be analysed with other molecular makers.
引文
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