中国佛蝗亚科（Phlaeobinae）的分类研究（直翅目：镌瓣亚目：蝗总科：剑角蝗科）
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摘要
佛蝗亚科Phlaeobinae隶属直翅目Orthoptera、镌瓣亚目Caelifera、蝗总科Acridoidea、剑角蝗科Acrididae，广泛分布于全世界。佛蝗亚科Phlaeobinae最早于1975年由DirshV. M.建立，对于该亚科分类系统的构建一直是蝗虫分类学的热点之一。本文对中国佛蝗亚科Phlaeobinae种类进行了采集、记录和描述，发现佛蝗属Phlaeoba1新种，并在印象初院士蝗虫进化理论的指导下建立了菊蝗亚科Phlaeobidinae subfamily nov.新亚科，该亚科的建立并得到分子分类学的印证。研究结果如下：
1、中国佛蝗亚科的分类
详细描述了中国佛蝗亚科、佛蝗类5个属的特征，编制了佛蝗亚科5个属的检索表和各属分种检索表，描述了佛蝗类23种的形态特征。发现并记述佛蝗属Phlaeoba南投佛蝗Phlaeoba nantouensis sp. nov.1新种（Ye and Yin,2007）。首次记述墨脱佛蝗Phlaeobamedogensis Liu,1981雌虫形态特征。补充记述《中国动物志·昆虫纲第三十二卷直翅目·蝗总科·槌角蝗科剑角蝗科》未记载的台湾佛蝗Phlaeoba formosana (Shiraki,1910)形态特征。
2、新亚科菊蝗亚科
陈世骧院士在《进化论和分类学》（科学出版社，1987）中提出在分类时要区分原始特征和新生特征。原直翅目Protorthoptera化石的翅很长，説明长翅是原始特征。印象初院士在《蝗虫类在青藏高原上的适应性》（1981）一文中提出蝗虫的演化途径是：长翅→短翅→无翅；发音器发达→退化→消失；鼓膜器（听器）发达→退化→消失；就是说缺翅、缺发音器、缺听器的种类是最进化的种类。该理论观点在中国蝗总科Acridoidea分类系统的研究、欧亚大陆斑翅蝗科分类系统研究中得到良好应用。另外2012年张道川等在《Insect Science》上发表1个缺翅、缺发音器、缺听器的新属：高原蝗属Pacris Zhanget al.，并建立一个新亚科Pacrinae高原蝗亚科，填补了棒角蝗科Gomphoceridae中最进化类群的空白。所有这一些都充分证明印象初院士蝗虫进化理论的正确性。
全世界剑角蝗类群中尚未发现缺翅、缺发音器、缺听器的种类，缺发音器的在传统佛蝗亚科Phlaeobinae中仅有2个属：菊蝗属（Phlaeobida Bolivar,1902）和华菊蝗属（Sinophlaeobida Yin et Yin,2007）。根据传统佛蝗亚科内菊蝗属Phlaeobida、华菊蝗属Sinophlaeobida的种类前翅短小、置于体两侧，不能发音，与其他佛蝗亚科属种类有明显区别，认为两个属的种类是比其他属较进化的类群。因此提出建立菊蝗亚科Phlaeobidinae subfamily nov.新亚科，模式属为菊蝗属Phlaeobida Bolivar,1902。编制了新亚科分属种检索表和剑角蝗科Acrididae分亚科检索表,描述了菊蝗类4个种的形态特征。
菊蝗亚科Phlaeobidinae subfamily nov.新亚科的设立也得到分子分类结果的支持。
在Gene Bank里选择了隆额网翅蝗Arcyptera coreana、李氏大足蝗Aeropus licenti、中华剑角蝗Acrida cinerea、白纹佛蝗Phlaeoba albonema之4种蝗虫的线粒体全基因组中cox2片段，结合自测的弱线菊蝗Phlaeobida carinata（PC）、海南菊蝗Phlaeobidahainanensis（PH）、暗色佛蝗Phlaeoba tenebrosa（PY采自友谊关，PR采自瑞丽)的cox2基因，利用软件进行比对并构建系统发育树。结果显示，菊蝗亚科Phlaeobidinae形成独立的进化分支，与剑角蝗亚科Acridinae组成姐妹群，再与佛蝗亚科Phlaeobinae组成姐妹群，而非菊蝗亚科Phlaeobidinae和佛蝗亚科Phlaeobinae直接组成姐妹群，说明了菊蝗亚科Phlaeobidinae与剑角蝗亚科Acridinae具有较近亲缘关系，与佛蝗亚科Phlaeobinae亲缘关系较远。
菊蝗亚科Phlaeobidinae subfam. nov.的建立，是以翅、发音器、听器特征为基础的蝗虫进化理论的指导下完成的，是剑角蝗科Acrididae分类的又一新研究成果，进一步丰富了蝗虫进化理论的内涵和剑角蝗科系统学研究。同时，新亚科的建立对于剑角蝗科系统学深入研究（如分子系统学方面）和在全世界范围发现菊蝗亚科物种多样性方面具有参考意义。
Phlaeobinae, a subfamily of Acrididae, belonging to Orthoptera, Caelifera, Acridoidea,distribute worldwide and was established by Dirsh V. M. in1975. Among which somemorphologically evolutionary taxa existed. The Systematics of the Phlaeobinae now remainssome arguments, and the establishment of its system is a hot spot in grasshopper systemicresearches. The taxa of Phlaeobinae from China was described in this thesis, and one newspecies, Phlaeoba nantouensis sp. nov. was found and described. Under the guide of Yin’grasshopper evolution theory, a new subfamily, Phlaeobidinae subfamily nov. in Acrididae,was established and further confirmed well with molecular data. All the results were as below.
1. Systematics of Phlaeobinae from China
The characteristic of the subfamily Phlaeobinae and the genus Phlaeoba were describedin detail, and the key to5genera and their species in the subfamily is given.23species inPhlaeobinae from China were described and illustrated. A new species Phlaeoba nantouensissp. nov. was found and described (Ye and Yin,2007). The morphological characteristics offemale of Phlaeoba medogensis Liu,1981was described for the first time. Another species,Phlaeoba formosana (Shiraki,1910), were also supplemented in the thesis, which were notrecorded in “Fauna sinica: Insect, Vol.32: Orthoptera: Acridoidea: Gomphoceridae andAcirididae”.
2. A new subfamily Phlaeobidinae subfam. nov. in Acrididae
In1987, Academician Chen showed that to distinguish characters into original orevolutionary when working taxonomy (“Evolutionary theory and systematics” SciencesPress). In fossil of Protorthoptera, having long tegmen, so the long tegmen is an originalcharacter. Yin (1981) put forward the evolutional pathways of grasshoppers: long tegmina→short tegmina→without tegmina, stridulatory organ developed→degenerated→disappearedand tympanal organ developed→degenerated→disappeared (“Adaptations of grasshoppersand locusts in Qinghai-Xizang Plateau”), so taxa lacking tegmina and wing, lackingstridulatory organ and lacking tympanal organ are most evolved. Later, this viewpoint wasapplied well in “On the taxonomic system of Acridoidea from China”(Yin,1982) and “On thetaxonomic system of Eurasian Oedipodidae”(Yin and Wang,2005). More recently, Zhang et al (2012) published a new grasshopper genus Pacris Zhang et al. without tegmina and wing,stridulatory organ and tympanal organ and established a new subfamily, Pacrinae is a mostevolved taxas in Gomphoceridae. So they are confirmed the evolutionary theory ofgrasshoppers,4species in Phlaeobidinae from China were described.
At present, the most evolved species without tegmina and wings, stridulatory organ andtympanal organ have not been found in Acrididae in the world. Based on short tegminapositioned laterally and without stridulatory organ in two genera, Phlaeobida Bolivar,1902and Sinophlaeobida Yin et Yin，2007, quite different from other genera in traditionalPhlaeobinae, we argued that the two genera are relative more evolutional taxa and establisheda new subfamily Phlaeobidinae in Acrididae. The type genus is Phlaeobida Bolivar,1902.The new subfamily includes Phlaeobida Bolivar,1902and Sinophlaeobida Yin et Yin,2007.Meanwhile, keys to genera in the new subfamily and subfamilies in Acrididae are given inthis paper.
The establishment of new subfamily was supported by molecular data. We chosen cox2gene of4species of grasshoppers, Arcyptera coreana, Aeropus licenti, Acrida cinerea, andPhlaeoba albonema in the their total genome sequences of mitochondrion in GenBank,together with cox2gene from Phlaeobida carinata（PC）, Phlaeobida hainanensis（PH）andPhlaeoba tenebrosa (PY) determined by ourselves, aligned these cox2gene sequences byusing bioinformatics softwares and established the molecular phylogenetic tree, whichshowed that the two species, Phlaeobida carinata and Phlaeobida hainanensis, in the newsubfamily was combined into a clade, generating a sister group with Acridinae, the sistergroup with Phlaeobinae composed another sister group. The phylogenetic tree also displayedthat Phlaeobidinae and Phlaeobinae do not compose a sister group directly, showingPhlaeobidinae has a closer relationship with Acridinae than Phlaeobinae.
The establishment of the new subfamily Phlaeobidinae，based on the evolutional theory ongrasshoppers according to the evolutional pathways of some morphological characters, is anew research achievement, which further enrich evolutional theory on grasshoppers andperfect the systematic researches in Acrididae. Meanwhile，it has important theoretic value forus to understand the phylogenetic relationship between different taxa and the evolutionalrelationship among different taxa in the family Acrididae, especially it will direct the grasshopper researchers to find more taxa in the world for increasing species diversity in thenew subfamily.
1、中国佛蝗亚科的分类
详细描述了中国佛蝗亚科、佛蝗类5个属的特征，编制了佛蝗亚科5个属的检索表和各属分种检索表，描述了佛蝗类23种的形态特征。发现并记述佛蝗属Phlaeoba南投佛蝗Phlaeoba nantouensis sp. nov.1新种（Ye and Yin,2007）。首次记述墨脱佛蝗Phlaeobamedogensis Liu,1981雌虫形态特征。补充记述《中国动物志·昆虫纲第三十二卷直翅目·蝗总科·槌角蝗科剑角蝗科》未记载的台湾佛蝗Phlaeoba formosana (Shiraki,1910)形态特征。
2、新亚科菊蝗亚科
陈世骧院士在《进化论和分类学》（科学出版社，1987）中提出在分类时要区分原始特征和新生特征。原直翅目Protorthoptera化石的翅很长，説明长翅是原始特征。印象初院士在《蝗虫类在青藏高原上的适应性》（1981）一文中提出蝗虫的演化途径是：长翅→短翅→无翅；发音器发达→退化→消失；鼓膜器（听器）发达→退化→消失；就是说缺翅、缺发音器、缺听器的种类是最进化的种类。该理论观点在中国蝗总科Acridoidea分类系统的研究、欧亚大陆斑翅蝗科分类系统研究中得到良好应用。另外2012年张道川等在《Insect Science》上发表1个缺翅、缺发音器、缺听器的新属：高原蝗属Pacris Zhanget al.，并建立一个新亚科Pacrinae高原蝗亚科，填补了棒角蝗科Gomphoceridae中最进化类群的空白。所有这一些都充分证明印象初院士蝗虫进化理论的正确性。
全世界剑角蝗类群中尚未发现缺翅、缺发音器、缺听器的种类，缺发音器的在传统佛蝗亚科Phlaeobinae中仅有2个属：菊蝗属（Phlaeobida Bolivar,1902）和华菊蝗属（Sinophlaeobida Yin et Yin,2007）。根据传统佛蝗亚科内菊蝗属Phlaeobida、华菊蝗属Sinophlaeobida的种类前翅短小、置于体两侧，不能发音，与其他佛蝗亚科属种类有明显区别，认为两个属的种类是比其他属较进化的类群。因此提出建立菊蝗亚科Phlaeobidinae subfamily nov.新亚科，模式属为菊蝗属Phlaeobida Bolivar,1902。编制了新亚科分属种检索表和剑角蝗科Acrididae分亚科检索表,描述了菊蝗类4个种的形态特征。
菊蝗亚科Phlaeobidinae subfamily nov.新亚科的设立也得到分子分类结果的支持。
在Gene Bank里选择了隆额网翅蝗Arcyptera coreana、李氏大足蝗Aeropus licenti、中华剑角蝗Acrida cinerea、白纹佛蝗Phlaeoba albonema之4种蝗虫的线粒体全基因组中cox2片段，结合自测的弱线菊蝗Phlaeobida carinata（PC）、海南菊蝗Phlaeobidahainanensis（PH）、暗色佛蝗Phlaeoba tenebrosa（PY采自友谊关，PR采自瑞丽)的cox2基因，利用软件进行比对并构建系统发育树。结果显示，菊蝗亚科Phlaeobidinae形成独立的进化分支，与剑角蝗亚科Acridinae组成姐妹群，再与佛蝗亚科Phlaeobinae组成姐妹群，而非菊蝗亚科Phlaeobidinae和佛蝗亚科Phlaeobinae直接组成姐妹群，说明了菊蝗亚科Phlaeobidinae与剑角蝗亚科Acridinae具有较近亲缘关系，与佛蝗亚科Phlaeobinae亲缘关系较远。
菊蝗亚科Phlaeobidinae subfam. nov.的建立，是以翅、发音器、听器特征为基础的蝗虫进化理论的指导下完成的，是剑角蝗科Acrididae分类的又一新研究成果，进一步丰富了蝗虫进化理论的内涵和剑角蝗科系统学研究。同时，新亚科的建立对于剑角蝗科系统学深入研究（如分子系统学方面）和在全世界范围发现菊蝗亚科物种多样性方面具有参考意义。
Phlaeobinae, a subfamily of Acrididae, belonging to Orthoptera, Caelifera, Acridoidea,distribute worldwide and was established by Dirsh V. M. in1975. Among which somemorphologically evolutionary taxa existed. The Systematics of the Phlaeobinae now remainssome arguments, and the establishment of its system is a hot spot in grasshopper systemicresearches. The taxa of Phlaeobinae from China was described in this thesis, and one newspecies, Phlaeoba nantouensis sp. nov. was found and described. Under the guide of Yin’grasshopper evolution theory, a new subfamily, Phlaeobidinae subfamily nov. in Acrididae,was established and further confirmed well with molecular data. All the results were as below.
1. Systematics of Phlaeobinae from China
The characteristic of the subfamily Phlaeobinae and the genus Phlaeoba were describedin detail, and the key to5genera and their species in the subfamily is given.23species inPhlaeobinae from China were described and illustrated. A new species Phlaeoba nantouensissp. nov. was found and described (Ye and Yin,2007). The morphological characteristics offemale of Phlaeoba medogensis Liu,1981was described for the first time. Another species,Phlaeoba formosana (Shiraki,1910), were also supplemented in the thesis, which were notrecorded in “Fauna sinica: Insect, Vol.32: Orthoptera: Acridoidea: Gomphoceridae andAcirididae”.
2. A new subfamily Phlaeobidinae subfam. nov. in Acrididae
In1987, Academician Chen showed that to distinguish characters into original orevolutionary when working taxonomy (“Evolutionary theory and systematics” SciencesPress). In fossil of Protorthoptera, having long tegmen, so the long tegmen is an originalcharacter. Yin (1981) put forward the evolutional pathways of grasshoppers: long tegmina→short tegmina→without tegmina, stridulatory organ developed→degenerated→disappearedand tympanal organ developed→degenerated→disappeared (“Adaptations of grasshoppersand locusts in Qinghai-Xizang Plateau”), so taxa lacking tegmina and wing, lackingstridulatory organ and lacking tympanal organ are most evolved. Later, this viewpoint wasapplied well in “On the taxonomic system of Acridoidea from China”(Yin,1982) and “On thetaxonomic system of Eurasian Oedipodidae”(Yin and Wang,2005). More recently, Zhang et al (2012) published a new grasshopper genus Pacris Zhang et al. without tegmina and wing,stridulatory organ and tympanal organ and established a new subfamily, Pacrinae is a mostevolved taxas in Gomphoceridae. So they are confirmed the evolutionary theory ofgrasshoppers,4species in Phlaeobidinae from China were described.
At present, the most evolved species without tegmina and wings, stridulatory organ andtympanal organ have not been found in Acrididae in the world. Based on short tegminapositioned laterally and without stridulatory organ in two genera, Phlaeobida Bolivar,1902and Sinophlaeobida Yin et Yin，2007, quite different from other genera in traditionalPhlaeobinae, we argued that the two genera are relative more evolutional taxa and establisheda new subfamily Phlaeobidinae in Acrididae. The type genus is Phlaeobida Bolivar,1902.The new subfamily includes Phlaeobida Bolivar,1902and Sinophlaeobida Yin et Yin,2007.Meanwhile, keys to genera in the new subfamily and subfamilies in Acrididae are given inthis paper.
The establishment of new subfamily was supported by molecular data. We chosen cox2gene of4species of grasshoppers, Arcyptera coreana, Aeropus licenti, Acrida cinerea, andPhlaeoba albonema in the their total genome sequences of mitochondrion in GenBank,together with cox2gene from Phlaeobida carinata（PC）, Phlaeobida hainanensis（PH）andPhlaeoba tenebrosa (PY) determined by ourselves, aligned these cox2gene sequences byusing bioinformatics softwares and established the molecular phylogenetic tree, whichshowed that the two species, Phlaeobida carinata and Phlaeobida hainanensis, in the newsubfamily was combined into a clade, generating a sister group with Acridinae, the sistergroup with Phlaeobinae composed another sister group. The phylogenetic tree also displayedthat Phlaeobidinae and Phlaeobinae do not compose a sister group directly, showingPhlaeobidinae has a closer relationship with Acridinae than Phlaeobinae.
The establishment of the new subfamily Phlaeobidinae，based on the evolutional theory ongrasshoppers according to the evolutional pathways of some morphological characters, is anew research achievement, which further enrich evolutional theory on grasshoppers andperfect the systematic researches in Acrididae. Meanwhile，it has important theoretic value forus to understand the phylogenetic relationship between different taxa and the evolutionalrelationship among different taxa in the family Acrididae, especially it will direct the grasshopper researchers to find more taxa in the world for increasing species diversity in thenew subfamily.
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