中国益蝽亚科修订及蠋蝽属、辉蝽属和二星蝽属的DNA分类学研究（半翅目：蝽科）
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摘要
本论文对中国益蝽亚科(半翅目：蝽科)进行了修订,并对蠋蝽属(益蝽亚科)、辉蝽属(蝽亚科)、二星蝽属(蝽亚科)在以往基于形态学数据修订的基础上进行DNA分类学研究。全文共分为三部分：
第一部分：主要总结了益蝽亚科的国内外研究历史与现状,DNA分类学的研究进展。详细介绍了本论文涉及的研究材料、研究方法,以及用于益蝽亚科分类的形态学特征及本文中使用的相应术语和量度。简单的介绍了益蝽亚科的生物学特性及经济意义。
第二部分：记述了中国益蝽亚科的修订结果。本文共记载中国益蝽亚科17属57种：其中包含6新种(其中2新种己发表),3新异名(其中1个已发表)1恢复异名以及1中国新纪录种。
新种：宽丹蝽Amyotea lata Zhao, Bu&Liu,2011；翘益蝽Picromerus elevatus Zhao, Liu&Bu,2013;红象蝽Cecyrina rubra sp. nov.;黄斑象蝽Cecyrina flavimacula sp. nov.;平象蝽Cecyrina flata sp. nov.;小象蝽Cecyrina parva sp. nov.o
新异名：膜疣蝽Cazira membrania Zhang&Lin,1982(=峨嵋疣蝽Cazira emeia Zhang&Lin,1982);四川疣蝽Cazira sichuana Zhang&Lin,1986(=削疣蝽Cazira frivaldszkyi Horvath,1889);红角并蝽Pinthaeus humeralis Horvath,1911(=并蝽Pinthaeus sanguinipes (Fabricius,1781))(已发表)。
恢复异名：蝎蝽Arma chinensis Fallou,1881恢复为欧亚蠋蝽Arma custos (Fabricius,1794)的异名;
中国新纪录种：宽曙厉蝽Eocanthecona shikokuensis (Esaki&Ishihara,1950)。
文中编制了中国益蝽亚科的属种检索表。对所研究种类进行了详细记述：包括检视了保存于国内的模式标本,并通过国际同行核对了部分保存于国外的模式标本；对重要分类特征进行重新描述；丰富了已知种的分布记录；对未见标本的种提供了原始描述。同时提供了成虫背腹面照片114幅,雌雄性生殖节照片88幅,还绘制了雄虫生殖节结构特征图43幅,以及提供每个分类单元的文献引证。
第三部分：提供了蠋蝽属(3种)、辉蝽属(6种)、二星蝽属(10种)DNA分类学研究结果。
本研究选取了中国蠋蝽属3种19头个体,辉蝽属6种50头个体以及二星蝽属10种70头个体作为研究对象,利用扩增的细胞色素氧化酶I (Cytochrome Coxidase subunit I)基因(约600bp)和细胞色素b (Cytochrome b)基因(蝎蝽属约410bp,辉蝽属约720bp)序列,以及GenBank中已有的一条序列,构建系统发育树、计算遗传距离,统计核苷酸多样性等。结果显示,蝎蝽属的种内遗传距离均小于0.02,蠋蝽和欧亚蝎蝽之间的遗传距离为0.0125,而朝鲜蝎蝽与蝎蝽和欧亚蝎蝽之间的遗传距离分别为0.1113和0.1079；以益蝽为外群进行的系统发育分析结果也不支持蝎蝽和欧亚蝎蝽各自的单系性；并经过对大量标本的比较研究发现这两个种间在阳基侧突的变异方面存在过渡类型,因此本文中恢复蝎蝽为欧亚蝎蝽的异名。辉蝽属各种的种内平均遗传距离都小于0.01；辉蝽与北方辉蝽之间的遗传距离为0.004,远小于其他种间平均遗传距离(0.075-0.15),属于种内遗传距离的范围；以二星蝽为外群的系统发育结果显示,虽然北方辉蝽的不同单倍型形成一个支系,但与辉蝽若干单倍型形成多分支关系,且辉蝽和北方辉蝽共同形成支系的支持率均很高(BI的PP为1.00或ML的BP为100)；在形态上,辉蝽和北方辉蝽外观几不可辨,生殖节中只有阳基侧突略有差异,但这种差异属于辉蝽种内的变异范围,因此,基于形态和分子数据建议将辉蝽和北方辉蝽合并为一个种,待查看模式标本后再做正式处理。二星蝽属的种内平均遗传距离差距较大,在0.003-0.039之间；而种间遗传距离在0.062-0.193之间；近缘种间遗传距离明显小于远缘种间遗传距离；系统发育结果显示,种间遗传距离较小的种间关系更近缘,但各自均能形成独立的支系；基于遗传距离和系统发育树的结果均支持Josifov&Kerzhner(1978)将尖角二星蝽作为北二星蝽异名的处理。本研究结果显示DNA分类学在解决属内种间的分类时可提供新的观点和新的视角,可结合形态与其他数据共同解决分类学问题。
新种的模式标本除注明外均保存于南开大学昆虫学研究所。
This dissertation deals with the taxonomic revision of the subfamily Asopinae (Hemiptera:Pentatomidae) from China and the DNA taxonomy of three genera, Arma (Asopinae), Carbula (Pentatominae) and Eysarcoris (Pentatominae), based on COI and cytb sequences. Three parts are included:introduction, taxonomic revision and DNA taxonomy.
The introduction briefly introduces the research history of Asopinae and DNA taxonomy, provides the descriptions of the general morphology and genital characters, the materials and methods used in the dissertation, abbreviations, measurements and the summary of biology and economic importance of Asopinae are also provided.
The taxonomy part treats with17genera57species of Asopinae from China.6species are described as new to the science.3new synonyms and1reinstate synonym are proposed, and1species is newly recorded to China.
New species:Amyotea lata Zhao, Bu&Liu,2011; Picromerus elevatus Zhao, Liu&Bu,2013; Cecyrina rubra sp. nov.; Cecyrina flavimacula sp. nov.; Cecyrina flata sp. nov.; Cecyrina parva sp. nov.
New synonyms:Cazira membrania Zhang&Lin,1982(=Cazira emeia Zhang&Lin,1982); Cazira sichuana Zhang&Lin,1986(=Cazira frivaldszkyi Horvath,1889); Pinthaeus humeralis Horvath,1911(=Pinthaeus sanguinipes (Fabricius,1781)).
Reinstate synonym:Arma chinensis Fallou,1881was recovered as a synonym of Arma custos (Fabricius,1794).
Chinese new record:Eocanthecona shikokuensis (Esaki&Ishihara,1950).
This dissertation offers the keys to the genera from China and to the species of each genus of Asopinae. Redescriptions of important characters, the updated distributional information, and the original descriptions of the unchecked species are provided. Most of the type specimens deposited in China were checked and we also checked some type specimens which deposited in other countries via the aid of colleagues. The detailed descriptions of species,43illustrations of male genitalia,144photographs of dorsal and ventral habitus,88photographs of female and male genitalia are provided. Citations for each taxon are listed.
The last part is the DNA taxonomy of three genera Arma, Carbula and Eysarcoris. The COI gene fragments (about600bp) and cytb gene fragments(Arma genus about410bp, Carbula genus about720bp) were amplified and sequenced. This research sampled19individuals of3species in Arma genus,50individuals of6species in genus Carbula and70individuals of10species in genus Eysarcoris and1sequence from GenBank as aimed objects, the data of gene COI and cytb were sequenced to construct phylogenetic tree, calculate genetic distance and compute nucleotide polymorphism. The result shows that all the genetic distance among the genus Arma are less than0.02, the genetic distance between Arma custos and Arma koreana, Arma chinensis and Arma koreana, is0.1113and0.1079respectively, but the distance between Arma custos and Arma chinensis is0.0125, which supports the latter two are the same species. Meanwhile, the phylogenetic tree construction also denied the monophyly of them respectively. So both of the morphological and molecular data of our research show a strong tendency to the reinstate synonym of Arma chinensis as Arma custos. The interspecific genetic distance among the genus Carbula is smaller than0.01, while the intraspecific ones ranges from0.075to0.15, besides0.004between Carbula humerigera and Carbula putoni. According to the average interspecific distance, the genetic distance between the two is more likely to be an interspecific one. The phylogenetic tree reveals that some haplotypes of C. putoni gathered together and paraphylied with the other haplotypes of C. humerigera. The monophyly of C. humerigera and C. putoni were strongly supported (the PP of BI is1.00and the BP of ML is100) in every tree. Morphologically, the appearence of C. humerigera and C. putoni can't be recognized from each other, the disecction of male genitalia shows little distinction in paramere, which can also be found intraspecifically, indicating that this character can't be used as diagnosis. So, based on the morphological and molecular data, we consider C. humerigera and C. putoni as the same species and will propose formally after checking the type specimens. The average interspecific genetic distance among the genus Eysarcoris varies from0.003 to0.039, while the intraspecific ones ranges from0.062to0.192. The intraspecific distance of sibling species is much smaller than that of distant species. Phylogenetic tree showed the monophyly of two species whose intraspecific distance is small. Besides, the outcome based on genetic distance and phylogenetic tree construction both support the opinion of Josifov&Kerzhner (1978), which proposed the synonym of Eysarcoris parvus. Consequently, the research showed DNA taxonomy can provide the new ideas and new perspective, and can be combined with morphology and other data to resolve the taxonomy.
The type specimens of new species are deposited in the Institute of Entomology, College of Life Science, Nankai University, Tianjin, China, except noted.
第一部分：主要总结了益蝽亚科的国内外研究历史与现状,DNA分类学的研究进展。详细介绍了本论文涉及的研究材料、研究方法,以及用于益蝽亚科分类的形态学特征及本文中使用的相应术语和量度。简单的介绍了益蝽亚科的生物学特性及经济意义。
第二部分：记述了中国益蝽亚科的修订结果。本文共记载中国益蝽亚科17属57种：其中包含6新种(其中2新种己发表),3新异名(其中1个已发表)1恢复异名以及1中国新纪录种。
新种：宽丹蝽Amyotea lata Zhao, Bu&Liu,2011；翘益蝽Picromerus elevatus Zhao, Liu&Bu,2013;红象蝽Cecyrina rubra sp. nov.;黄斑象蝽Cecyrina flavimacula sp. nov.;平象蝽Cecyrina flata sp. nov.;小象蝽Cecyrina parva sp. nov.o
新异名：膜疣蝽Cazira membrania Zhang&Lin,1982(=峨嵋疣蝽Cazira emeia Zhang&Lin,1982);四川疣蝽Cazira sichuana Zhang&Lin,1986(=削疣蝽Cazira frivaldszkyi Horvath,1889);红角并蝽Pinthaeus humeralis Horvath,1911(=并蝽Pinthaeus sanguinipes (Fabricius,1781))(已发表)。
恢复异名：蝎蝽Arma chinensis Fallou,1881恢复为欧亚蠋蝽Arma custos (Fabricius,1794)的异名;
中国新纪录种：宽曙厉蝽Eocanthecona shikokuensis (Esaki&Ishihara,1950)。
文中编制了中国益蝽亚科的属种检索表。对所研究种类进行了详细记述：包括检视了保存于国内的模式标本,并通过国际同行核对了部分保存于国外的模式标本；对重要分类特征进行重新描述；丰富了已知种的分布记录；对未见标本的种提供了原始描述。同时提供了成虫背腹面照片114幅,雌雄性生殖节照片88幅,还绘制了雄虫生殖节结构特征图43幅,以及提供每个分类单元的文献引证。
第三部分：提供了蠋蝽属(3种)、辉蝽属(6种)、二星蝽属(10种)DNA分类学研究结果。
本研究选取了中国蠋蝽属3种19头个体,辉蝽属6种50头个体以及二星蝽属10种70头个体作为研究对象,利用扩增的细胞色素氧化酶I (Cytochrome Coxidase subunit I)基因(约600bp)和细胞色素b (Cytochrome b)基因(蝎蝽属约410bp,辉蝽属约720bp)序列,以及GenBank中已有的一条序列,构建系统发育树、计算遗传距离,统计核苷酸多样性等。结果显示,蝎蝽属的种内遗传距离均小于0.02,蠋蝽和欧亚蝎蝽之间的遗传距离为0.0125,而朝鲜蝎蝽与蝎蝽和欧亚蝎蝽之间的遗传距离分别为0.1113和0.1079；以益蝽为外群进行的系统发育分析结果也不支持蝎蝽和欧亚蝎蝽各自的单系性；并经过对大量标本的比较研究发现这两个种间在阳基侧突的变异方面存在过渡类型,因此本文中恢复蝎蝽为欧亚蝎蝽的异名。辉蝽属各种的种内平均遗传距离都小于0.01；辉蝽与北方辉蝽之间的遗传距离为0.004,远小于其他种间平均遗传距离(0.075-0.15),属于种内遗传距离的范围；以二星蝽为外群的系统发育结果显示,虽然北方辉蝽的不同单倍型形成一个支系,但与辉蝽若干单倍型形成多分支关系,且辉蝽和北方辉蝽共同形成支系的支持率均很高(BI的PP为1.00或ML的BP为100)；在形态上,辉蝽和北方辉蝽外观几不可辨,生殖节中只有阳基侧突略有差异,但这种差异属于辉蝽种内的变异范围,因此,基于形态和分子数据建议将辉蝽和北方辉蝽合并为一个种,待查看模式标本后再做正式处理。二星蝽属的种内平均遗传距离差距较大,在0.003-0.039之间；而种间遗传距离在0.062-0.193之间；近缘种间遗传距离明显小于远缘种间遗传距离；系统发育结果显示,种间遗传距离较小的种间关系更近缘,但各自均能形成独立的支系；基于遗传距离和系统发育树的结果均支持Josifov&Kerzhner(1978)将尖角二星蝽作为北二星蝽异名的处理。本研究结果显示DNA分类学在解决属内种间的分类时可提供新的观点和新的视角,可结合形态与其他数据共同解决分类学问题。
新种的模式标本除注明外均保存于南开大学昆虫学研究所。
This dissertation deals with the taxonomic revision of the subfamily Asopinae (Hemiptera:Pentatomidae) from China and the DNA taxonomy of three genera, Arma (Asopinae), Carbula (Pentatominae) and Eysarcoris (Pentatominae), based on COI and cytb sequences. Three parts are included:introduction, taxonomic revision and DNA taxonomy.
The introduction briefly introduces the research history of Asopinae and DNA taxonomy, provides the descriptions of the general morphology and genital characters, the materials and methods used in the dissertation, abbreviations, measurements and the summary of biology and economic importance of Asopinae are also provided.
The taxonomy part treats with17genera57species of Asopinae from China.6species are described as new to the science.3new synonyms and1reinstate synonym are proposed, and1species is newly recorded to China.
New species:Amyotea lata Zhao, Bu&Liu,2011; Picromerus elevatus Zhao, Liu&Bu,2013; Cecyrina rubra sp. nov.; Cecyrina flavimacula sp. nov.; Cecyrina flata sp. nov.; Cecyrina parva sp. nov.
New synonyms:Cazira membrania Zhang&Lin,1982(=Cazira emeia Zhang&Lin,1982); Cazira sichuana Zhang&Lin,1986(=Cazira frivaldszkyi Horvath,1889); Pinthaeus humeralis Horvath,1911(=Pinthaeus sanguinipes (Fabricius,1781)).
Reinstate synonym:Arma chinensis Fallou,1881was recovered as a synonym of Arma custos (Fabricius,1794).
Chinese new record:Eocanthecona shikokuensis (Esaki&Ishihara,1950).
This dissertation offers the keys to the genera from China and to the species of each genus of Asopinae. Redescriptions of important characters, the updated distributional information, and the original descriptions of the unchecked species are provided. Most of the type specimens deposited in China were checked and we also checked some type specimens which deposited in other countries via the aid of colleagues. The detailed descriptions of species,43illustrations of male genitalia,144photographs of dorsal and ventral habitus,88photographs of female and male genitalia are provided. Citations for each taxon are listed.
The last part is the DNA taxonomy of three genera Arma, Carbula and Eysarcoris. The COI gene fragments (about600bp) and cytb gene fragments(Arma genus about410bp, Carbula genus about720bp) were amplified and sequenced. This research sampled19individuals of3species in Arma genus,50individuals of6species in genus Carbula and70individuals of10species in genus Eysarcoris and1sequence from GenBank as aimed objects, the data of gene COI and cytb were sequenced to construct phylogenetic tree, calculate genetic distance and compute nucleotide polymorphism. The result shows that all the genetic distance among the genus Arma are less than0.02, the genetic distance between Arma custos and Arma koreana, Arma chinensis and Arma koreana, is0.1113and0.1079respectively, but the distance between Arma custos and Arma chinensis is0.0125, which supports the latter two are the same species. Meanwhile, the phylogenetic tree construction also denied the monophyly of them respectively. So both of the morphological and molecular data of our research show a strong tendency to the reinstate synonym of Arma chinensis as Arma custos. The interspecific genetic distance among the genus Carbula is smaller than0.01, while the intraspecific ones ranges from0.075to0.15, besides0.004between Carbula humerigera and Carbula putoni. According to the average interspecific distance, the genetic distance between the two is more likely to be an interspecific one. The phylogenetic tree reveals that some haplotypes of C. putoni gathered together and paraphylied with the other haplotypes of C. humerigera. The monophyly of C. humerigera and C. putoni were strongly supported (the PP of BI is1.00and the BP of ML is100) in every tree. Morphologically, the appearence of C. humerigera and C. putoni can't be recognized from each other, the disecction of male genitalia shows little distinction in paramere, which can also be found intraspecifically, indicating that this character can't be used as diagnosis. So, based on the morphological and molecular data, we consider C. humerigera and C. putoni as the same species and will propose formally after checking the type specimens. The average interspecific genetic distance among the genus Eysarcoris varies from0.003 to0.039, while the intraspecific ones ranges from0.062to0.192. The intraspecific distance of sibling species is much smaller than that of distant species. Phylogenetic tree showed the monophyly of two species whose intraspecific distance is small. Besides, the outcome based on genetic distance and phylogenetic tree construction both support the opinion of Josifov&Kerzhner (1978), which proposed the synonym of Eysarcoris parvus. Consequently, the research showed DNA taxonomy can provide the new ideas and new perspective, and can be combined with morphology and other data to resolve the taxonomy.
The type specimens of new species are deposited in the Institute of Entomology, College of Life Science, Nankai University, Tianjin, China, except noted.
引文
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