中国锯天牛族系统发育分析
摘要
锯天牛族(Prionini)隶属于鞘翅目(Coleoptera)天牛科(Cerambycidae)锯天牛亚科(Prioninae),其中的不少种类是农林业生产中的重要害虫。对锯天牛族昆虫的研究以传统的外部形态为主,并且均为种类的分类和鉴定,到目前为止还没有关于锯天牛族线粒体基因的研究报道,锯天牛族各分类单元之间的系统发育关系也不清楚,因此本研究对中国锯天牛族的线粒体12S rRNA、16S rRNA、COI基因进行了研究,并依据传统的外部形态特征、后翅翅脉和雄性外生殖器特征、线粒体基因序列分析了中国锯天牛族的系统发育关系,具体结果如下：
DNA提取及线粒体部分基因扩增
馆藏干标本DNA发生了明显降解,降解随着标本保存时间的增加而增加。电泳检测发现从保存二十年干标本提取的DNA扩增到了目的条带,但测序结果却表明这些片段并不是天牛本身的基因片段。单纯采用PCR技术和琼脂糖凝胶电泳技术作为评价DNA提取方法优劣的指标并不可靠。本研究共得到中国锯天牛族3属8种和寡刺细齿天牛的mtDNA 12S rRNA、16S rRNA和COI基因序列24条。
线粒体基因序列分析
本研究发现,12S rRNA和16S rRNA基因发生了碱基插入(缺失),COI基因则没有。研究得到的3个线粒体基因序列的碱基组成均明显偏向于T和A,核糖体RNA基因更偏向于A,而COI基因稍偏向于T。12S rRNA基因的碱基转换／颠换率随着分化时间的增加而减小,16S rRNA基因先增加后减小,COI基因则不发生明显变化。COI基因的进化速度快于12S rRNA基因和16S rRNA基因,其进化速度主要体现在第三密码子位点上,第二密码子位点的进化速度很慢。以不同基因序列计算的物种之间遗传距离的大小关系存在差异。并且依据线粒体基因序列计算的P距离小于实际的遗传距离。
沟翅土天牛Dorysthenes fossatus (Pascoe,1857) 12S rRNA基因序列长度,12SrRNA、16S rRNA和12S rRNA+16S rRNA基因序列的碱基组成和土天牛属其他种类存在差异。沟翅土天牛与土天牛属Dorysthenes Vigors,1826其他种类之间的遗传距离和碱基转换／颠换率明显不同于土天牛属其他种类之间的遗传距离和碱基转换／颠换率,与锯天牛属Prionus Geoffroy,1762和土天牛属之间的遗传距离和碱基转换／颠换率近似。
中国锯天牛族的系统发育关系分析
邻接搜索法得到的最小距离树并不一定是真正的最小距离树,利用调整后保留指数对各特征进行加权可以优化简约法的分析结果。系统发育一致树和最优树之间的拓扑结构并不完全相同,有的甚至出现相互矛盾的分支,而进化模型对最小距离树拓扑结构的影响大于距离一致树。不同系统发育分析方法的分析结果并不一定相同,但是似然法的分析结果在Bayes法中均有体现。依据不同特征重建的系统发育关系可能存在冲突。
以寡刺细齿天牛Rhaphipodus fruhstorferi Lameere,1903和黄星天牛Psacothea hilaris (Pascoe,1857)为外群,发现中国锯天牛族是一个单系群,应该分为两个亚族,锯天牛属和土天牛属归入Prioni亚族,接眼天牛属Priotyrannus Thomson,1857和拟土天牛属Prionomma White,1853归入Derobrachi亚族。
中国锯天牛属是一个多系群。缪氏锯天牛Prionus murzini Drumont et Komiya,2006、单叶锯天牛Prionus unilamellatus Pu,1987和赛氏锯天牛Prionus siskai Drumont et Komiya,2006应该从锯天牛属中独立出来成为一个单独的属,其中单叶锯天牛和赛氏锯天牛之间的亲缘关系更近。短角锯天牛Prionus gahani Lameere,1912、锯天牛Prionus insular is Motschulsky,1857、齿跗锯天牛Prionus sifanicus Plavilstshikov,1934、皱胸锯天牛罗氏亚种Prionus delavayi lorenci Drumont et Komiya,2006和云南锯天牛Prionus lameerei Semenov-Tian-Shanskij,1927组成一个单系群,其中短角锯天牛、齿跗锯天牛和云南锯天牛之间的亲缘关系更近。
中国土天牛属是一个多系群。Dorysthenes (Prionomimus)亚属应该从土天牛属中分离出去成为一个独立的属,其中丽土天牛Dorysthenes elegans Ohbayashi,1981和沟翅土天牛之间的亲缘关系更近。除去Dorysthenes (Prionomimus)亚属之后,土天牛属的种类组成一个单系群,证明Lameere依据外部形态特征将Lophosternus Guerin,1844, Paraphrus Thomson,1860, Cyrtognathus Faldermann,1835和Baladeva Waterhouse,1840等4属并入土天牛属成为新的土天牛属的做法是合理的。蔗根土天牛Dorysthenes granulosus (Thomson,1860)是土天牛属中较原始的类群。Dorysthenes (Cyrtognathus)亚属是一个单系群,其中大牙土天牛Dorysthenes paradoxus (Faldermann,1833)和甘肃土天牛Dorysthenes tippmanni Heyrovsky,1950具有更近的亲缘关系。依据外部形态特征建立的Dorysthenes(Lophosternus)亚属和Dorysthenes (Baladeva)亚属的种类并没有分别聚成区别于其他种类的分支。
Tribe Prionini is one of the oldest tribes in Prioninae (Coleoptera:Cerambycidae) and many species in the tribe are important pests in agriculture and forestry. Studies on Prionini were focused on morphology, especially on identification and taxonomy. Up to now, mitochondrial genes of Prionini are not studied systematicly, and phylogenetic relationships among taxa in Prionini are unknown. The dissertation analysed partial 12S rRNA,16S rRNA and COI gene sequences and discussed the phylogenetic relationship among taxa in Chinese Prionini based on traditional morphological characters, characters of hind wing venation and male genetalia, and mitochondrial gene sequences. Results are followed:
DNA extraction and PCR amplification of three mitochondrial genes
In dry specimens, apparent DNA degration was found and long deposition time accelerated DNA degration. After PCR amplification based on DNA from dry specimens deposited longer than 20 years, mitochondrial gene segments were found by agarose gel electrophoresis technique. However, sequences of these mitochondrial gene segments were different from those of Cerambycidae. In the study,24 gene sequences of mtDNA 12S rRNA,16S rRNA and COI genes were obteined from 8 species of 3 genera in Chinese Prionini and a species of Macrotomini.
Analysis of mitochondrial gene sequences in Prionini
Base insertion/miss was found in 12S rRNA and 16S rRNA genes, but not in COI gene. Base constitution of obtained mithchondrial genes biased towards T and A. In ribosomal rRNA gene, content of base A was higher than base T, but content of base T higher than base A in COI gene. Ratios of transition vs transversion (Ts/Tv) in 12S rRNA gene decreased when the divergence time increased. Ts/Tv ratios in 16S rRNA gene first increased but then decreased when the divergence time increased. Ts/Tv ratios in COI gene sequences did not changed distinctly when the divergence time increased. The evolution rate of COI gene was faster than ribosomal rRNA genes and the evolution in COI gene was primarily located in the third coden sites. The evolution rates of these second coden sites were very slow. Evolution distances based on diverse genes were different. P evolution distances were smaller than these true evolution distances.
Dorysthenes fossatus (Pascoe,1857) was different from other species of Dorysthenes in the length of 12S rRNA gene segment and base constitutions of 12S rRNA,16S rRNA and 12S rRNA+16S rRNA gene sequences. The evolution distances and Ts/Tv ratios between Dorysthenes fossatus and other species of Dorysthenes were different from those among species of Dorysthenes (excluding Dorysthenes fossatus), but is the same as those between Dorysthenes and Prionus.
Phylogenetic analysis of Chinese Prionini
The evolution scores of these best distance trees given by Neighbor joint method were usually not the minimum scores. Weighting characters by rescaled consistency index (RC) was useful for maximum parsimonious analysis. The topology of bootstrap 50% majority-rule consensus tree was different from the best phylogenetic tree usually. The best phylogenetic tree was more influenced by substution model than bootstrap 50% majority-rule consensus tree. The topologies of phylogenetic trees reconstructed from different methods may be different. However the topology of phylogenetic trees reconstructed from maximum likehood method appeared in phylogenetic trees reconstructed from Bayesian method. The topologies of phylogenetic trees reconstructed from different characters may be not same.
Chinese Prionini is a monophyletic group and should be divided into two subtribes. Genera Dorysthenes and Prionus belongs to subtribe Prioni, and genera Priotyrannus and Prionomma belongs to subtribe Derobrachi.
Genus Prionus defined by morphological characters of mandible is paraphyletic. Prionus murzini Drumont et Komiya,2006, Prionus unilamellatus Pu,1987 and Prionus siskai Drumont et Komiya,2006 should be excluded from genus Prionus and constitute a new independent genus. There is close relationship between Prionus unilamellatus and Prionus siskai. Prionus gahani Lameere,1912, Prionus insularis Motschulsky,1857, Prionus sifanicus Plavilstshikov,1934, Prionus delavayi lorenci Drumont et Komiya,2006 and Prionus lameerei Semenov-Tian-Shanskij,1927 constitute a monophyletic group. There are close relationship among Prionus gahani, Prionus lameerei and Prionus sifanicus.
Genus Dorysthenes defined by morphological characters of mandible is a paraphyletic group too. Subgenus Dorysthenes (Prionomimus) should be excluded from genus Dorysthenes and constitute a new genus. There is close relationship between Dorysthenes elegans Ohbayashi,1981 and Dorysthenes fossatus. Genus Dorysthenes is a monophological group after excluding subgenus Dorysthenes (Prionomimus). It is reasonable that Lameere combined genera Lophosternus Guerin, 1844, Paraphrus Thomson,1860, Cyrtognathus Faldermann,1835 and Baladeva Waterhouse,1840 with genus Dorysthenes based on morphological characters. Dorysthenes granulosus (Thomson,1860) is the primitive species of genus Dorysthenes. Subgenus Dorysthenes(Cyrtognathus) is monophological, and there is close relationship between Dorysthenes paradoxus (Faldermann,1833) and Dorysthenes tippmanni Heyrovsky,1950. Subgenera Dorysthenes (Lophosternus) and Dorysthenes (Baladeva) defined by traditional morphological characters are paraphyletic.
DNA提取及线粒体部分基因扩增
馆藏干标本DNA发生了明显降解,降解随着标本保存时间的增加而增加。电泳检测发现从保存二十年干标本提取的DNA扩增到了目的条带,但测序结果却表明这些片段并不是天牛本身的基因片段。单纯采用PCR技术和琼脂糖凝胶电泳技术作为评价DNA提取方法优劣的指标并不可靠。本研究共得到中国锯天牛族3属8种和寡刺细齿天牛的mtDNA 12S rRNA、16S rRNA和COI基因序列24条。
线粒体基因序列分析
本研究发现,12S rRNA和16S rRNA基因发生了碱基插入(缺失),COI基因则没有。研究得到的3个线粒体基因序列的碱基组成均明显偏向于T和A,核糖体RNA基因更偏向于A,而COI基因稍偏向于T。12S rRNA基因的碱基转换／颠换率随着分化时间的增加而减小,16S rRNA基因先增加后减小,COI基因则不发生明显变化。COI基因的进化速度快于12S rRNA基因和16S rRNA基因,其进化速度主要体现在第三密码子位点上,第二密码子位点的进化速度很慢。以不同基因序列计算的物种之间遗传距离的大小关系存在差异。并且依据线粒体基因序列计算的P距离小于实际的遗传距离。
沟翅土天牛Dorysthenes fossatus (Pascoe,1857) 12S rRNA基因序列长度,12SrRNA、16S rRNA和12S rRNA+16S rRNA基因序列的碱基组成和土天牛属其他种类存在差异。沟翅土天牛与土天牛属Dorysthenes Vigors,1826其他种类之间的遗传距离和碱基转换／颠换率明显不同于土天牛属其他种类之间的遗传距离和碱基转换／颠换率,与锯天牛属Prionus Geoffroy,1762和土天牛属之间的遗传距离和碱基转换／颠换率近似。
中国锯天牛族的系统发育关系分析
邻接搜索法得到的最小距离树并不一定是真正的最小距离树,利用调整后保留指数对各特征进行加权可以优化简约法的分析结果。系统发育一致树和最优树之间的拓扑结构并不完全相同,有的甚至出现相互矛盾的分支,而进化模型对最小距离树拓扑结构的影响大于距离一致树。不同系统发育分析方法的分析结果并不一定相同,但是似然法的分析结果在Bayes法中均有体现。依据不同特征重建的系统发育关系可能存在冲突。
以寡刺细齿天牛Rhaphipodus fruhstorferi Lameere,1903和黄星天牛Psacothea hilaris (Pascoe,1857)为外群,发现中国锯天牛族是一个单系群,应该分为两个亚族,锯天牛属和土天牛属归入Prioni亚族,接眼天牛属Priotyrannus Thomson,1857和拟土天牛属Prionomma White,1853归入Derobrachi亚族。
中国锯天牛属是一个多系群。缪氏锯天牛Prionus murzini Drumont et Komiya,2006、单叶锯天牛Prionus unilamellatus Pu,1987和赛氏锯天牛Prionus siskai Drumont et Komiya,2006应该从锯天牛属中独立出来成为一个单独的属,其中单叶锯天牛和赛氏锯天牛之间的亲缘关系更近。短角锯天牛Prionus gahani Lameere,1912、锯天牛Prionus insular is Motschulsky,1857、齿跗锯天牛Prionus sifanicus Plavilstshikov,1934、皱胸锯天牛罗氏亚种Prionus delavayi lorenci Drumont et Komiya,2006和云南锯天牛Prionus lameerei Semenov-Tian-Shanskij,1927组成一个单系群,其中短角锯天牛、齿跗锯天牛和云南锯天牛之间的亲缘关系更近。
中国土天牛属是一个多系群。Dorysthenes (Prionomimus)亚属应该从土天牛属中分离出去成为一个独立的属,其中丽土天牛Dorysthenes elegans Ohbayashi,1981和沟翅土天牛之间的亲缘关系更近。除去Dorysthenes (Prionomimus)亚属之后,土天牛属的种类组成一个单系群,证明Lameere依据外部形态特征将Lophosternus Guerin,1844, Paraphrus Thomson,1860, Cyrtognathus Faldermann,1835和Baladeva Waterhouse,1840等4属并入土天牛属成为新的土天牛属的做法是合理的。蔗根土天牛Dorysthenes granulosus (Thomson,1860)是土天牛属中较原始的类群。Dorysthenes (Cyrtognathus)亚属是一个单系群,其中大牙土天牛Dorysthenes paradoxus (Faldermann,1833)和甘肃土天牛Dorysthenes tippmanni Heyrovsky,1950具有更近的亲缘关系。依据外部形态特征建立的Dorysthenes(Lophosternus)亚属和Dorysthenes (Baladeva)亚属的种类并没有分别聚成区别于其他种类的分支。
Tribe Prionini is one of the oldest tribes in Prioninae (Coleoptera:Cerambycidae) and many species in the tribe are important pests in agriculture and forestry. Studies on Prionini were focused on morphology, especially on identification and taxonomy. Up to now, mitochondrial genes of Prionini are not studied systematicly, and phylogenetic relationships among taxa in Prionini are unknown. The dissertation analysed partial 12S rRNA,16S rRNA and COI gene sequences and discussed the phylogenetic relationship among taxa in Chinese Prionini based on traditional morphological characters, characters of hind wing venation and male genetalia, and mitochondrial gene sequences. Results are followed:
DNA extraction and PCR amplification of three mitochondrial genes
In dry specimens, apparent DNA degration was found and long deposition time accelerated DNA degration. After PCR amplification based on DNA from dry specimens deposited longer than 20 years, mitochondrial gene segments were found by agarose gel electrophoresis technique. However, sequences of these mitochondrial gene segments were different from those of Cerambycidae. In the study,24 gene sequences of mtDNA 12S rRNA,16S rRNA and COI genes were obteined from 8 species of 3 genera in Chinese Prionini and a species of Macrotomini.
Analysis of mitochondrial gene sequences in Prionini
Base insertion/miss was found in 12S rRNA and 16S rRNA genes, but not in COI gene. Base constitution of obtained mithchondrial genes biased towards T and A. In ribosomal rRNA gene, content of base A was higher than base T, but content of base T higher than base A in COI gene. Ratios of transition vs transversion (Ts/Tv) in 12S rRNA gene decreased when the divergence time increased. Ts/Tv ratios in 16S rRNA gene first increased but then decreased when the divergence time increased. Ts/Tv ratios in COI gene sequences did not changed distinctly when the divergence time increased. The evolution rate of COI gene was faster than ribosomal rRNA genes and the evolution in COI gene was primarily located in the third coden sites. The evolution rates of these second coden sites were very slow. Evolution distances based on diverse genes were different. P evolution distances were smaller than these true evolution distances.
Dorysthenes fossatus (Pascoe,1857) was different from other species of Dorysthenes in the length of 12S rRNA gene segment and base constitutions of 12S rRNA,16S rRNA and 12S rRNA+16S rRNA gene sequences. The evolution distances and Ts/Tv ratios between Dorysthenes fossatus and other species of Dorysthenes were different from those among species of Dorysthenes (excluding Dorysthenes fossatus), but is the same as those between Dorysthenes and Prionus.
Phylogenetic analysis of Chinese Prionini
The evolution scores of these best distance trees given by Neighbor joint method were usually not the minimum scores. Weighting characters by rescaled consistency index (RC) was useful for maximum parsimonious analysis. The topology of bootstrap 50% majority-rule consensus tree was different from the best phylogenetic tree usually. The best phylogenetic tree was more influenced by substution model than bootstrap 50% majority-rule consensus tree. The topologies of phylogenetic trees reconstructed from different methods may be different. However the topology of phylogenetic trees reconstructed from maximum likehood method appeared in phylogenetic trees reconstructed from Bayesian method. The topologies of phylogenetic trees reconstructed from different characters may be not same.
Chinese Prionini is a monophyletic group and should be divided into two subtribes. Genera Dorysthenes and Prionus belongs to subtribe Prioni, and genera Priotyrannus and Prionomma belongs to subtribe Derobrachi.
Genus Prionus defined by morphological characters of mandible is paraphyletic. Prionus murzini Drumont et Komiya,2006, Prionus unilamellatus Pu,1987 and Prionus siskai Drumont et Komiya,2006 should be excluded from genus Prionus and constitute a new independent genus. There is close relationship between Prionus unilamellatus and Prionus siskai. Prionus gahani Lameere,1912, Prionus insularis Motschulsky,1857, Prionus sifanicus Plavilstshikov,1934, Prionus delavayi lorenci Drumont et Komiya,2006 and Prionus lameerei Semenov-Tian-Shanskij,1927 constitute a monophyletic group. There are close relationship among Prionus gahani, Prionus lameerei and Prionus sifanicus.
Genus Dorysthenes defined by morphological characters of mandible is a paraphyletic group too. Subgenus Dorysthenes (Prionomimus) should be excluded from genus Dorysthenes and constitute a new genus. There is close relationship between Dorysthenes elegans Ohbayashi,1981 and Dorysthenes fossatus. Genus Dorysthenes is a monophological group after excluding subgenus Dorysthenes (Prionomimus). It is reasonable that Lameere combined genera Lophosternus Guerin, 1844, Paraphrus Thomson,1860, Cyrtognathus Faldermann,1835 and Baladeva Waterhouse,1840 with genus Dorysthenes based on morphological characters. Dorysthenes granulosus (Thomson,1860) is the primitive species of genus Dorysthenes. Subgenus Dorysthenes(Cyrtognathus) is monophological, and there is close relationship between Dorysthenes paradoxus (Faldermann,1833) and Dorysthenes tippmanni Heyrovsky,1950. Subgenera Dorysthenes (Lophosternus) and Dorysthenes (Baladeva) defined by traditional morphological characters are paraphyletic.
引文
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