中国苔蚁甲族的分类学及属级系统发育研究
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摘要
苔蚁甲族(Tyrini)隶属于鞘翅目(Coleoptera),隐翅虫科(Staphylinidae),蚁甲亚科(Pselaphinae),是长角蚁甲超族(Pselaphitae)下的一个大族。该族成员形态相似,体型中至大型,广泛存在于农、林生态系统中,全部为捕食性昆虫,是食物链和食物网的重要一环,对生态平衡起着重要的作用,是自然界不可或缺的成员。
本研究之前,全世界苔蚁甲族已知81属601种,分布于世界各大动物区系,其中以新热带区、澳洲区及东洋区较为集中。我国苔蚁甲族已知14属42种,仅占全世界已知种类的7.0%。其中,东洋区成分占主导,成员较为集中地分布于华东、华南及西南各省份山区。国内对苔蚁甲族的分类研究起步较晚,研究基础相对薄弱,主要有两方面原因：其一,大部分已知种由国外学者零星记载,模式标本流落国外。尤其是发表于18世纪末到19世纪初的不少种类,其描述简单,没有涉及雄性外生殖器等重要的分类学特征,且不含插图,很多种自原描述后再未有过记录,因此对这些物种的鉴定存在较大的困难。其二,国内许多生物多样性热点地区从未利用现代的采集技术对苔蚁甲族资源进行过系统的调查,导致我国该类群生物多样性和资源本底的情况仍严重不清,妨碍了后续开发和利用。因此,对我国苔蚁甲族资源进行深入调查、开展系统分类研究是非常必要和迫切的。
博士学习期间,作者采用经典形态分类学和支序系统学的原理和方法,通过检视多年来采自我国70多个自然保护区的3,500余号苔蚁甲族标本,核查大量保存于大英自然博物馆、瑞士自然历史博物馆、日本国立自然博物馆、捷克国家博物馆和保加利亚国家自然历史博物馆等国外知名博物馆的馆藏模式标本,并借阅了德国、捷克、俄罗斯等国际同行学者私人收藏的数百号未定名标本,对我国苔蚁甲族种类进行了系统研究,结果如下：
一、共记载我国苔蚁甲族15属94种(本研究前为14属42种),论文包含14属91种,其中发现新属1属,新种52种,即：大瑶蚁甲属Dayaogen. nov.、彭氏瑶蚁甲Dayao pengzhongi sp. nov.(广西)、峨眉瑶蚁甲Dayao emeiensis sp. nov.(四川)、版纳拟蚁甲Labomimus bannaus sp. nov.(云南)、邻拟蚁甲Labomimus cognatus sp. nov.(云南)、大巴山拟蚁甲Labomimus dabashanus sp. nov.(陕西、湖北)、毛拟蚁甲Labomimus fimbriatus sp. nov.(云南)、九顶山拟蚁甲Labomimus jiudingensis sp. nov.(四川)、鸡足山拟蚁甲Labomimus jizuensis sp. nov.(云南)、猫儿山拟蚁甲Labomimus maoershanus sp. nov.(广西)、奇拟蚁甲Labomimus mirus sp. nov.(云南)、类突拟蚁甲Labomimus paratorus sp. nov.(陕西)、方胸拟蚁甲Labomimus quadratithorax sp. nov.(广西)、耙拟蚁甲Labomimus sarculus sp. nov.(云南)、舒克氏拟蚁甲Labomimus schuelkei sp. nov.(陕西)、须拟蚁甲Labomimus simplicipalpus sp. nov.(云南)、胫拟蚁甲Labomimus tibialis sp. nov.(云南)、鸡足山拟蚁甲Labomimus venustus sp. nov.(云南)、蝠拟蚁甲Labomimus vespertilio sp. nov.(云南)、东方毛蚁甲Lasinus orientalis sp. nov.(浙江)、幸运安蚁甲Linanfortunatus sp. nov.(广西）、花坪安蚁甲Linan huapingensis sp. nov.(广西)、胡氏安蚁甲hujiayaoi sp. nov.(广西)、胸安蚁甲Linan tendothorax sp. nov.(云南)、上野氏安蚁甲Linan wewoi sp. nov.(广西)、舒克氏巨苔蚁甲Wegatyrus schuelkei sp. nov.(云南)、腾冲巨苔蚁甲Megatyrus tengchongensis sp. nov.(云南)、中华野村蚁甲Nomuraius sinicus sp. nov.(广西)、安徽长角蚁甲Pselaphodes anhuianus sp. nov.(安徽)、戴氏长角蚁甲Pselaphodes daii sp. nov.(四川)、大瑶山长角蚁甲Pselaphodes dayaoensis sp.nov.(广西)、奇茎长角蚁甲Pselaphodes distincticornis sp. nov.(云南)、二郎山长角蚁甲Pselaphodes erlangshanus sp. nov.(四川)、曲角长角蚁甲Pselaphodes flexus sp. nov.(云南)、格雷氏长角蚁甲Pselaphodes grebennikovi sp. nov.(云南)、海南长角蚁甲Pselaphodes hainanensis sp. nov.(海南)、胡氏长角蚁甲Pselaphodes hui sp. nov.(广西)、岸本氏长角蚁甲Pselaphodes kishimotoi sp. nov.(贵州)、宽阔水长角蚁甲Pselaphodes kuankuoshuiensis sp. nov.(贵州)、长茎长角蚁甲Pselaphodes longilobus sp. nov.(云南)、猫儿山长角蚁甲Pselaphodes maoershanus sp. nov.(广西)、独角长角蚁甲Pselaphodes monoceros sp. nov.(西藏)、林氏长角蚁甲Pselaphodes linae sp.nov.(福建、海南)、彭氏长角蚁甲Pselaphodes pengi sp. nov.(四川)、拟沃克氏长角蚁甲Pselaphodes pseudowalkeri sp. nov.(浙江、江西、福建)、史氏长角蚁甲Pselaphodes shii sp. nov.(海南)、天童长角蚁甲Pselaphodes tiantongensis sp. nov.(浙江)、莱士长角蚁甲Pselaphodes wrasei sp. nov.(云南)、中甸长角蚁甲Pselaphodes zhongdianus sp. nov.(云南)、野村氏鼻苔蚁甲Tyrinasius nomurai sp. nov.(宁夏)、杰氏拟苔蚁甲Tyrodes jenisi sp. nov.(云南)、四川苔蚁甲Tyrus sichuanicus sp. nov.(四川)及雅江苔蚁甲Tyrus yajiangensis sp. nov.(四川)(上述新种均已另文正式发表)。对于我国已记载的种类,大部分通过赴国外博物馆借阅、核查模式标本或赴模式产地采集地模标本等途径与手段,进行了形态学的比较研究和再描述。
二、给出了所有涉及种类详的细描述、研究历史、采集数据和分布地区,讨论了各物种的鉴定特征以及与近缘种的区分要点。所有种类均配有彩色整体形态图,其中89种附有解剖图版。
三、编制了中国苔蚁甲族分属、分种组和分种检索表。
四、整理了中国苔蚁甲族名录。
五、利用FileMaker Pro Advanced11.0v3软件,构建了全世界苔蚁甲族所有已知属、种的数据库,其中包含物种名、定名人、定名年代、模式产地、分布、研究历史等相关分类学信息,从而在宏观上实现对该类群的区系构成、地理分布及研究历史的快速统计与分析。
六、采用支序系统学的方法,以我国苔蚁甲族28个代表种作为内群,奇须蚁甲族(Tmesiphorini)下属的隆背蚁甲属(Saltisedes)2种作为外群,选取57个性状状态,对其进行解释和编码。利用WinClada ver.1.00.08和NONA ver.2.0软件,对我国苔蚁甲族下2亚族13属的系统发育关系进行了初步分析。结果表明,苔蚁甲族下长角蚁甲属团(Pselaphodes complex of genera)包含的中国类群拥有极高的单系支持度,在进化树上聚为一支,拥有较近的亲缘关系,符合现今的分类系统。但作为锥须蚁甲亚族(Centrophthalmina)成员的锥须蚁甲属Centrophthalmus却落在苔蚁甲亚族(Tyrina)内部,这有两种可能,一是由于分析采用非加性特征,所有性状不分权重,导致分析结果偏离实际情况。二是传统分类系统过度强调下颚须形态的重要性,而人为地将锥须蚁甲属及3个近缘属归入单独的亚族。
Members of the beetle tribe Tyrini (Coleoptera, Staphylinidae, Pselaphinae, Pselaphitae) are medium to large in size, morphologically similar predators commonly found in forest and farmland ecosystems. They are believed to act as an essential link in food chain and food web, and have significant importance of ecological equilibrium.
Before this study,603species scattered in81genera of the Tyrini have been known worldwide, with the Neotropical, Australian, and Oriental regions have the highest species diversity. Fourteen genera and42species have been known to occur in China, accounting for7.0%of the total world fauna. While having a dominance of Oriental elements over the country, most species are distributed in mountain areas of eastern, southern, and southwestern provinces. Taxonomic studies on the tribe Tyrini by domestic scientists started relatively late in China. Many taxa need a modern revision and an easy-to-use identification key. This is mostly caused by the follo wings:1) Type material of many known species described by foreign workers, are housed in respective foreign museums. Some 'old'species have never been reported again since their original description between the late18th and the first half of the19th century. The simple descriptions of these species, and the lack of illustrations caused much difficulties in species identification.2) Proper samplings using modern techniques have never been conducted in many biodiversity hotspots in China, even estimated numbers of species from these areas remain unavailable, hindering further development and utilization of this resource. Therefore, a large-scale investigation and a through taxonomic study on the Chinese species of Tyrini is necessary and in urgent need.
During the present study, over3,500specimens collected from more than70nature reserves in China, many types housed in the Natural History Museum, London, the Natural History Museum of Geneva, the National Museum of Nature and Science, Tokyo, the National Museum, Prague, Czech Republic, and the National Natural History Museum, Bulgarica, and hundreds of unidentified tyrine specimens from several private collections, are examined. The study revealed the following results:
1. A total of15genera and94species are now known in China, among which,14genera and91species are covered here. This includes one new genus, viz., Dayao gen. nov., and the following52new species:Dayao pengzhongi sp. nov.(Guangxi), Dayao emeiensis sp. nov.(Sichuan), Labomimus bannaus sp. nov.(Yunnan), Labomimus cognatus sp. nov.(Yunnan), Labomimus dabashanus sp. nov.(Shaanxi, Hubei), Labomimus fimbriatus sp. nov.(Yunnan), Labomimus jiudingensis sp. nov.(Sichuan), Labomimus jizuensis sp. nov.(Yunnan), Labomimus maoershanus sp. nov.(Guangxi), Labomimus mirus sp. nov.(Yunnan), Labomimus paratorus sp. nov.(Shaanxi), Labomimus quadratithorax sp. nov.(Guangxi), Labomimus sarculus sp. nov.(Yunnan), Labomimus schuelkei sp. nov.(Shaanxi), Labomimus simplicipalpus sp. nov.(Yunnan), Labomimus tibialis sp. nov.(Yunnan), Labomimus venustus sp. nov.(Yunnan), Labomimus vespertilio sp. nov.(Yunnan), Lasinus orientalis sp. nov.(Zhejiang), Linan fortunatus sp. nov.(Guangxi), Linan huapingensis sp. nov.(Guangxi), Linan hujiayaoi sp. nov.(Guangxi), Linan tendothorax sp. nov.(Yunnan), Linan uenoi sp. nov.(Guangxi), Megatyrus schuelkei sp. nov.(Yunnan), Megatyrus tengchongensis, sp. nov.(Yunnan), Nomuraius sinicus sp. nov.(Guangxi), Pselaphodes anhuianus sp. nov.(Anhui), Pselaphodes daii sp. nov.(Sichuan), Pselaphodes dayaoensis sp. nov.(Guangxi), Pselaphodes distincticornis sp. nov.(Yunnan), Pselaphodes erlangshanus sp. nov.(Sichuan), Pselaphodes flexus sp. nov.(Yunnan), Pselaphodes grebennikovi sp. nov.(Yunnan), Pselaphodes hainanensis sp. nov.(Hainan), Pselaphodes hui sp. nov.(Guangxi), Pselaphodes kishimotoi sp. nov.(Guizhou), Pselaphodes kuankuoshuiensis sp. nov.(Guizhou), Pselaphodes longilobus sp. nov.(Yunnan), Pselaphodes maoershanus sp. nov.(Guangxi), Pselaphodes monoceros sp. nov.(Xizang), Pselaphodes linae sp. nov.(Fujian, Hainan), Pselaphodes pengi sp. nov.(Sichuan), Pselaphodes pseudowalkeri sp. nov.(Zhejiang, Jiangxi, Fujian), Pselaphodes shii sp. nov.(Hainan), Pselaphodes tiantongensis sp. nov.(Zhejiang), Pselaphodes wrasei sp. nov.(Yunnan), Pselaphodes zhongdianus sp. nov.(Yunnan), Tyrinasius nomurai sp. nov.(Ningxia), Tyrodes jenisi sp. nov.(Yunnan), Tyrus sichuanicus sp. nov.(Sichuan), and Tyrus yajiangensis sp. nov.(Sichuan). All new taxa have been formally published during the two-year time of this study. Identifications of the previously known Chinese species are clarified based on the type series deposited in respective museums, or additional material collected from type localities of respective species.
2. All species treated in this paper are described in detail. A taxonomic history, collecting data, distributional information, and comparative notes are provided for each species. Illustrations of habitus for all species, and diagnostic characters for89species are given.
3. Keys to genera, species groups, and species of each genus are presented.
4. A list of the tyrine species known to occur in China is provided.
5. A database containing the following taxonomic information for all known species in the world is built, facilitating quick statistical analyses:scientific name, author, year of publication, type locality, distribution, and taxonomic history.
6. A preliminary phylogenetic study on the tyrine genera from China is conducted by using WinClada ver.1.00.08and NONA ver.2.0. The analysis was run for28in-group and2out-group taxa, based on57character states which are coded and explained. The result showed that the Chinese members of the Pselaphodes complex of genera is well supported as a monophyletic group, which agrees well with the current taxonomic system, while the genus Centrophthalmus belonging to the subtribe Centrophthalmina is found to locate within Tyrina. This might be resulted from:1) all non-additive characters were treated equally-weighted, or,2) the classification on the subtribe level, which is largely based on the form of maxillary palpi, is artificial.
本研究之前,全世界苔蚁甲族已知81属601种,分布于世界各大动物区系,其中以新热带区、澳洲区及东洋区较为集中。我国苔蚁甲族已知14属42种,仅占全世界已知种类的7.0%。其中,东洋区成分占主导,成员较为集中地分布于华东、华南及西南各省份山区。国内对苔蚁甲族的分类研究起步较晚,研究基础相对薄弱,主要有两方面原因：其一,大部分已知种由国外学者零星记载,模式标本流落国外。尤其是发表于18世纪末到19世纪初的不少种类,其描述简单,没有涉及雄性外生殖器等重要的分类学特征,且不含插图,很多种自原描述后再未有过记录,因此对这些物种的鉴定存在较大的困难。其二,国内许多生物多样性热点地区从未利用现代的采集技术对苔蚁甲族资源进行过系统的调查,导致我国该类群生物多样性和资源本底的情况仍严重不清,妨碍了后续开发和利用。因此,对我国苔蚁甲族资源进行深入调查、开展系统分类研究是非常必要和迫切的。
博士学习期间,作者采用经典形态分类学和支序系统学的原理和方法,通过检视多年来采自我国70多个自然保护区的3,500余号苔蚁甲族标本,核查大量保存于大英自然博物馆、瑞士自然历史博物馆、日本国立自然博物馆、捷克国家博物馆和保加利亚国家自然历史博物馆等国外知名博物馆的馆藏模式标本,并借阅了德国、捷克、俄罗斯等国际同行学者私人收藏的数百号未定名标本,对我国苔蚁甲族种类进行了系统研究,结果如下：
一、共记载我国苔蚁甲族15属94种(本研究前为14属42种),论文包含14属91种,其中发现新属1属,新种52种,即：大瑶蚁甲属Dayaogen. nov.、彭氏瑶蚁甲Dayao pengzhongi sp. nov.(广西)、峨眉瑶蚁甲Dayao emeiensis sp. nov.(四川)、版纳拟蚁甲Labomimus bannaus sp. nov.(云南)、邻拟蚁甲Labomimus cognatus sp. nov.(云南)、大巴山拟蚁甲Labomimus dabashanus sp. nov.(陕西、湖北)、毛拟蚁甲Labomimus fimbriatus sp. nov.(云南)、九顶山拟蚁甲Labomimus jiudingensis sp. nov.(四川)、鸡足山拟蚁甲Labomimus jizuensis sp. nov.(云南)、猫儿山拟蚁甲Labomimus maoershanus sp. nov.(广西)、奇拟蚁甲Labomimus mirus sp. nov.(云南)、类突拟蚁甲Labomimus paratorus sp. nov.(陕西)、方胸拟蚁甲Labomimus quadratithorax sp. nov.(广西)、耙拟蚁甲Labomimus sarculus sp. nov.(云南)、舒克氏拟蚁甲Labomimus schuelkei sp. nov.(陕西)、须拟蚁甲Labomimus simplicipalpus sp. nov.(云南)、胫拟蚁甲Labomimus tibialis sp. nov.(云南)、鸡足山拟蚁甲Labomimus venustus sp. nov.(云南)、蝠拟蚁甲Labomimus vespertilio sp. nov.(云南)、东方毛蚁甲Lasinus orientalis sp. nov.(浙江)、幸运安蚁甲Linanfortunatus sp. nov.(广西）、花坪安蚁甲Linan huapingensis sp. nov.(广西)、胡氏安蚁甲hujiayaoi sp. nov.(广西)、胸安蚁甲Linan tendothorax sp. nov.(云南)、上野氏安蚁甲Linan wewoi sp. nov.(广西)、舒克氏巨苔蚁甲Wegatyrus schuelkei sp. nov.(云南)、腾冲巨苔蚁甲Megatyrus tengchongensis sp. nov.(云南)、中华野村蚁甲Nomuraius sinicus sp. nov.(广西)、安徽长角蚁甲Pselaphodes anhuianus sp. nov.(安徽)、戴氏长角蚁甲Pselaphodes daii sp. nov.(四川)、大瑶山长角蚁甲Pselaphodes dayaoensis sp.nov.(广西)、奇茎长角蚁甲Pselaphodes distincticornis sp. nov.(云南)、二郎山长角蚁甲Pselaphodes erlangshanus sp. nov.(四川)、曲角长角蚁甲Pselaphodes flexus sp. nov.(云南)、格雷氏长角蚁甲Pselaphodes grebennikovi sp. nov.(云南)、海南长角蚁甲Pselaphodes hainanensis sp. nov.(海南)、胡氏长角蚁甲Pselaphodes hui sp. nov.(广西)、岸本氏长角蚁甲Pselaphodes kishimotoi sp. nov.(贵州)、宽阔水长角蚁甲Pselaphodes kuankuoshuiensis sp. nov.(贵州)、长茎长角蚁甲Pselaphodes longilobus sp. nov.(云南)、猫儿山长角蚁甲Pselaphodes maoershanus sp. nov.(广西)、独角长角蚁甲Pselaphodes monoceros sp. nov.(西藏)、林氏长角蚁甲Pselaphodes linae sp.nov.(福建、海南)、彭氏长角蚁甲Pselaphodes pengi sp. nov.(四川)、拟沃克氏长角蚁甲Pselaphodes pseudowalkeri sp. nov.(浙江、江西、福建)、史氏长角蚁甲Pselaphodes shii sp. nov.(海南)、天童长角蚁甲Pselaphodes tiantongensis sp. nov.(浙江)、莱士长角蚁甲Pselaphodes wrasei sp. nov.(云南)、中甸长角蚁甲Pselaphodes zhongdianus sp. nov.(云南)、野村氏鼻苔蚁甲Tyrinasius nomurai sp. nov.(宁夏)、杰氏拟苔蚁甲Tyrodes jenisi sp. nov.(云南)、四川苔蚁甲Tyrus sichuanicus sp. nov.(四川)及雅江苔蚁甲Tyrus yajiangensis sp. nov.(四川)(上述新种均已另文正式发表)。对于我国已记载的种类,大部分通过赴国外博物馆借阅、核查模式标本或赴模式产地采集地模标本等途径与手段,进行了形态学的比较研究和再描述。
二、给出了所有涉及种类详的细描述、研究历史、采集数据和分布地区,讨论了各物种的鉴定特征以及与近缘种的区分要点。所有种类均配有彩色整体形态图,其中89种附有解剖图版。
三、编制了中国苔蚁甲族分属、分种组和分种检索表。
四、整理了中国苔蚁甲族名录。
五、利用FileMaker Pro Advanced11.0v3软件,构建了全世界苔蚁甲族所有已知属、种的数据库,其中包含物种名、定名人、定名年代、模式产地、分布、研究历史等相关分类学信息,从而在宏观上实现对该类群的区系构成、地理分布及研究历史的快速统计与分析。
六、采用支序系统学的方法,以我国苔蚁甲族28个代表种作为内群,奇须蚁甲族(Tmesiphorini)下属的隆背蚁甲属(Saltisedes)2种作为外群,选取57个性状状态,对其进行解释和编码。利用WinClada ver.1.00.08和NONA ver.2.0软件,对我国苔蚁甲族下2亚族13属的系统发育关系进行了初步分析。结果表明,苔蚁甲族下长角蚁甲属团(Pselaphodes complex of genera)包含的中国类群拥有极高的单系支持度,在进化树上聚为一支,拥有较近的亲缘关系,符合现今的分类系统。但作为锥须蚁甲亚族(Centrophthalmina)成员的锥须蚁甲属Centrophthalmus却落在苔蚁甲亚族(Tyrina)内部,这有两种可能,一是由于分析采用非加性特征,所有性状不分权重,导致分析结果偏离实际情况。二是传统分类系统过度强调下颚须形态的重要性,而人为地将锥须蚁甲属及3个近缘属归入单独的亚族。
Members of the beetle tribe Tyrini (Coleoptera, Staphylinidae, Pselaphinae, Pselaphitae) are medium to large in size, morphologically similar predators commonly found in forest and farmland ecosystems. They are believed to act as an essential link in food chain and food web, and have significant importance of ecological equilibrium.
Before this study,603species scattered in81genera of the Tyrini have been known worldwide, with the Neotropical, Australian, and Oriental regions have the highest species diversity. Fourteen genera and42species have been known to occur in China, accounting for7.0%of the total world fauna. While having a dominance of Oriental elements over the country, most species are distributed in mountain areas of eastern, southern, and southwestern provinces. Taxonomic studies on the tribe Tyrini by domestic scientists started relatively late in China. Many taxa need a modern revision and an easy-to-use identification key. This is mostly caused by the follo wings:1) Type material of many known species described by foreign workers, are housed in respective foreign museums. Some 'old'species have never been reported again since their original description between the late18th and the first half of the19th century. The simple descriptions of these species, and the lack of illustrations caused much difficulties in species identification.2) Proper samplings using modern techniques have never been conducted in many biodiversity hotspots in China, even estimated numbers of species from these areas remain unavailable, hindering further development and utilization of this resource. Therefore, a large-scale investigation and a through taxonomic study on the Chinese species of Tyrini is necessary and in urgent need.
During the present study, over3,500specimens collected from more than70nature reserves in China, many types housed in the Natural History Museum, London, the Natural History Museum of Geneva, the National Museum of Nature and Science, Tokyo, the National Museum, Prague, Czech Republic, and the National Natural History Museum, Bulgarica, and hundreds of unidentified tyrine specimens from several private collections, are examined. The study revealed the following results:
1. A total of15genera and94species are now known in China, among which,14genera and91species are covered here. This includes one new genus, viz., Dayao gen. nov., and the following52new species:Dayao pengzhongi sp. nov.(Guangxi), Dayao emeiensis sp. nov.(Sichuan), Labomimus bannaus sp. nov.(Yunnan), Labomimus cognatus sp. nov.(Yunnan), Labomimus dabashanus sp. nov.(Shaanxi, Hubei), Labomimus fimbriatus sp. nov.(Yunnan), Labomimus jiudingensis sp. nov.(Sichuan), Labomimus jizuensis sp. nov.(Yunnan), Labomimus maoershanus sp. nov.(Guangxi), Labomimus mirus sp. nov.(Yunnan), Labomimus paratorus sp. nov.(Shaanxi), Labomimus quadratithorax sp. nov.(Guangxi), Labomimus sarculus sp. nov.(Yunnan), Labomimus schuelkei sp. nov.(Shaanxi), Labomimus simplicipalpus sp. nov.(Yunnan), Labomimus tibialis sp. nov.(Yunnan), Labomimus venustus sp. nov.(Yunnan), Labomimus vespertilio sp. nov.(Yunnan), Lasinus orientalis sp. nov.(Zhejiang), Linan fortunatus sp. nov.(Guangxi), Linan huapingensis sp. nov.(Guangxi), Linan hujiayaoi sp. nov.(Guangxi), Linan tendothorax sp. nov.(Yunnan), Linan uenoi sp. nov.(Guangxi), Megatyrus schuelkei sp. nov.(Yunnan), Megatyrus tengchongensis, sp. nov.(Yunnan), Nomuraius sinicus sp. nov.(Guangxi), Pselaphodes anhuianus sp. nov.(Anhui), Pselaphodes daii sp. nov.(Sichuan), Pselaphodes dayaoensis sp. nov.(Guangxi), Pselaphodes distincticornis sp. nov.(Yunnan), Pselaphodes erlangshanus sp. nov.(Sichuan), Pselaphodes flexus sp. nov.(Yunnan), Pselaphodes grebennikovi sp. nov.(Yunnan), Pselaphodes hainanensis sp. nov.(Hainan), Pselaphodes hui sp. nov.(Guangxi), Pselaphodes kishimotoi sp. nov.(Guizhou), Pselaphodes kuankuoshuiensis sp. nov.(Guizhou), Pselaphodes longilobus sp. nov.(Yunnan), Pselaphodes maoershanus sp. nov.(Guangxi), Pselaphodes monoceros sp. nov.(Xizang), Pselaphodes linae sp. nov.(Fujian, Hainan), Pselaphodes pengi sp. nov.(Sichuan), Pselaphodes pseudowalkeri sp. nov.(Zhejiang, Jiangxi, Fujian), Pselaphodes shii sp. nov.(Hainan), Pselaphodes tiantongensis sp. nov.(Zhejiang), Pselaphodes wrasei sp. nov.(Yunnan), Pselaphodes zhongdianus sp. nov.(Yunnan), Tyrinasius nomurai sp. nov.(Ningxia), Tyrodes jenisi sp. nov.(Yunnan), Tyrus sichuanicus sp. nov.(Sichuan), and Tyrus yajiangensis sp. nov.(Sichuan). All new taxa have been formally published during the two-year time of this study. Identifications of the previously known Chinese species are clarified based on the type series deposited in respective museums, or additional material collected from type localities of respective species.
2. All species treated in this paper are described in detail. A taxonomic history, collecting data, distributional information, and comparative notes are provided for each species. Illustrations of habitus for all species, and diagnostic characters for89species are given.
3. Keys to genera, species groups, and species of each genus are presented.
4. A list of the tyrine species known to occur in China is provided.
5. A database containing the following taxonomic information for all known species in the world is built, facilitating quick statistical analyses:scientific name, author, year of publication, type locality, distribution, and taxonomic history.
6. A preliminary phylogenetic study on the tyrine genera from China is conducted by using WinClada ver.1.00.08and NONA ver.2.0. The analysis was run for28in-group and2out-group taxa, based on57character states which are coded and explained. The result showed that the Chinese members of the Pselaphodes complex of genera is well supported as a monophyletic group, which agrees well with the current taxonomic system, while the genus Centrophthalmus belonging to the subtribe Centrophthalmina is found to locate within Tyrina. This might be resulted from:1) all non-additive characters were treated equally-weighted, or,2) the classification on the subtribe level, which is largely based on the form of maxillary palpi, is artificial.
引文
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Besuchet, C. Pselaphides recoltes au Nepal par le Prof. H. Janetschek (Coleoptera)[J]. Ber. naturw.-med. Ver. Innsbruck.1971,58 (1970), 313-318.
Besuchet, C. Synonymes et homonyme nouveaux de quelques genres de Pselaphides (Coleoptera)[J]. Revue Suisse de Zoologie.1986,93 (1), 257-264.
Besuchet, C. Pselaphides palearctiques nouveaux ou meconnus (Coleoptera: Staphylinidae:Pselaphinae)[J]. Revue Suisse de Zoologie.1999,106 (4), 789-811.
Brendel, E. The Pselaphidae of North America. A monograph[J]. In:Brendel E. & Wickham H.F. Bulletin of the Laboratory of Natural History, State University of Iowa.1890,1,216-304, pls 6-9.
Blattny, C. Revise Pselaphidu sbirky Helferovy Nar. Musea v Praze. Revision der Pselaphiden der Collection Heifer im National-Museum in Prag[J]. Sbornik Entomologickeho Oddeleni Narodniho Musea v Praze.1925,3, 179-222.
Casey, T.L. On a new genus of South African Pselaphidae[J]. Transactions of the Entomological Society of London.1887,381-383.
Castellini, G. Pselafidi afrotropicali:note geonomiche e descrizione di nuove specie (Coleoptera)[J]. Atti del Museo di Storia Naturale della Maremma. 1997,16,163-207.
Fairmaire, L. M. H Coleopteres d'Algerie rapportes par M. Ogier de Baulny[J]. Annales de la Societe Entomologique de France (4e ser.).1863,3, 637-648.
Hlavac, P. A Taxonomic Revision of the Tyrini of the Oriental Region. II. Systematic study on the genus Pselaphodes and its allied genera (Coleoptera:Staphylinidae:Pselaphinae) [J]. Annales de la Societe Entomologique de France[J].2003a,38(3),283-297.
Hlavac, P. A Taxonomic Revision of Tyrini of the Oriental Region. IV. Synonymy and redescription of Labomimus heterocerus (Coleoptera: Staphylinidae:Pselaphinae). Entomological Problems[J].2003b,33(1-2), 93-94.
Hlavac, P. & Chandler, D. S. World Catalogue of the species of Tyrini with a key to the genera (Coleoptera:Staphylinidae:Pselaphinae)[J]. Folia Heyrovskyana.2005,13(3),81-143
Hlavac, P., & Nomura, S. A Taxonomic Revision of Tyrini of the Oriental Region I. Paralasinus (Coleoptera:Staphylinidae:Pselaphinae), a new genus of Tyrina from Indochina[J]. Elytra.2001,29(1),163-174.
Hlavac, P., & Nomura, S. A taxonomic revision of Tyrini of the Oriental region III. Megatyrus (Coleoptera, Staphylinidae, Pselaphinae), a new genus of the Tyrina from China and Vietnam[J]. Elytra.2003,31(1),165-174.
Hlavac, P., Nomura, S. & Zhou, H.Z. Two new species of the genus Labomimus from China (Coleoptera:Staphylinidae, Pselaphinae) [J]. Species diversity. 2000,5 (2),149-153.
Jeannel, R. Les Pselaphides de I'Afrique orientale[J]. Memoires du Museum National d'Histoire Naturelle, (Zool.).1949,29(1),1-226.
Jeannel, R. Faune du Congo belge et du Ruanda-Urundi. II. Pselaphidae[J]. Annales du Musee du Congo Belge, (ser.8vo).1950,2 (1949),1-278.
Jeannel, R. Sur les Pselaphides (Coleoptera) de l'Inde septentrionale[J]. Bulletin of the British Museum (Natural History) (Entomology).1960,9,403-456.
Kurbatov, S.A. Le nouveau genre de Pselaphidae (Coleoptera) de l'Asie Centrale[J]. Russian Entomological Journal.1993,2 (1),51-52.
Lobl, I. Eine neue Art der Gattung Lasinus Sharp aus China (Col., Pselaphidae) [J]. Acta Rerum Naturalium Musei Nationalis Slovenici, Bratislava.1964, 10,45-48.
Lucas, R. Catalogus alphabeticus generum et subgenerum Coleopterorum orbis terrarum totius (famil., trib., subtr., sect. incl.)[J]. Pars I. Arch. Naturgesch., (A).1920,84 (1918), i-xxxi+1-696.
Motschulsky, V. de. Enumeration des nouvelles especes de Coleopteres rapportes par M. Victor Motschoulsky de son dernier voyage[J]. Bulletin de la Societe Imperiale des Naturalistes de Moscou.1851,24 (lere partie), 479-511.
Newton, A. F. & Chandler, D. S. World catalog of the genera of Pselaphidae (Coleoptera)[J]. Fieldiana, Zoology, new series.1989,53, i-iii+1-93.
Nomura, S. Taxonomical notes on Chinese species of the tyrine genus Tyrinasius (Coleoptera, Staphylinidae, Pselaphinae)[J]. Elytra.1999,27 (2), 485-498.
Panzer, G. W. F. Pselaphus mucronatus. Fauna insectorum Germaniae initia oder Deuschlands Insecten[M].1801, Heft 89, No.11.
Raffray, A. Etude sur les Pselaphides. V. Tableaux synoptiques. Notes et synonymie[J]. Revue d'Entomologie.1890a,9,81-172.
Raffray, A. Etude sur les Pselaphides. VI. Diagnoses des especes nouvelles sur lesquelles sont fondes des genres nouveaux[J]. Revue d'Entomologie[J]. 1890b,9,193-219, pls 2,3.
Raffray, A. Revision des Pselaphides des iles de Singapore et de Penang (Suite et fin) [J]. Revue d'Entomologie.1895,14,21-82, pl.2.
Raffray, A. Pselaphides nouveaux de Ceylan[J]. Annales de la Societe Entomologique de France.1901,70,27-30.
Raffray, A. Genera et catalogue des Pselaphides [J]. Annales de la Societe Entomologique de France.1904,73,1-400.
Raffray, A. Genera et catalogue des Pselaphides[J]. Annales de la Societe Entomologique de France.1905,73 (1904),401-476, pls 1-3.
Raffray, A. Especes nouvelles de Pselaphides exotiques (Coleopt.) [J]. Annales de la Societe Entomologique de France.1912,80 (1911),425-450.
Raffray, A. H. Sauter's Formosa-Ausbeute. Pselaphidae (Col.) II[J]. Supplementa Entomologica.1914,3,1-5.
Reitter, E. Versuch einer systematischen Eintheilung der Clavigeriden und Pselaphiden[J]. Verhandlungen der Naturforschenden Vereines in Brunn. 1882,20,177-211.
Schaufuss, L.W. Ueber Pselaphidengattungen[J]. Nunquam Otiosus.1877,2, 450-460.
Schmidt-Goebel, H.M. Beytrag zur einer Monographie der Pselaphen, enthaltend neue Species aus Asien[M]. Erste Lieferung. Gottlieb Haase Sohne, Prague.1838,1-16.,2 pls.
Sharp, D. Descriptions of new genera and species of Pselaphidae and Scydmaenidae from Australia and New Zealand[J]. Transactions of the Entomological Society of London.1874(4),483-518.
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Besuchet, C. Pselaphides recoltes au Nepal par le Prof. H. Janetschek (Coleoptera)[J]. Ber. naturw.-med. Ver. Innsbruck.1971,58 (1970), 313-318.
Besuchet, C. Synonymes et homonyme nouveaux de quelques genres de Pselaphides (Coleoptera)[J]. Revue Suisse de Zoologie.1986,93 (1), 257-264.
Besuchet, C. Pselaphides palearctiques nouveaux ou meconnus (Coleoptera: Staphylinidae:Pselaphinae)[J]. Revue Suisse de Zoologie.1999,106 (4), 789-811.
Brendel, E. The Pselaphidae of North America. A monograph[J]. In:Brendel E. & Wickham H.F. Bulletin of the Laboratory of Natural History, State University of Iowa.1890,1,216-304, pls 6-9.
Blattny, C. Revise Pselaphidu sbirky Helferovy Nar. Musea v Praze. Revision der Pselaphiden der Collection Heifer im National-Museum in Prag[J]. Sbornik Entomologickeho Oddeleni Narodniho Musea v Praze.1925,3, 179-222.
Casey, T.L. On a new genus of South African Pselaphidae[J]. Transactions of the Entomological Society of London.1887,381-383.
Castellini, G. Pselafidi afrotropicali:note geonomiche e descrizione di nuove specie (Coleoptera)[J]. Atti del Museo di Storia Naturale della Maremma. 1997,16,163-207.
Fairmaire, L. M. H Coleopteres d'Algerie rapportes par M. Ogier de Baulny[J]. Annales de la Societe Entomologique de France (4e ser.).1863,3, 637-648.
Hlavac, P. A Taxonomic Revision of the Tyrini of the Oriental Region. II. Systematic study on the genus Pselaphodes and its allied genera (Coleoptera:Staphylinidae:Pselaphinae) [J]. Annales de la Societe Entomologique de France[J].2003a,38(3),283-297.
Hlavac, P. A Taxonomic Revision of Tyrini of the Oriental Region. IV. Synonymy and redescription of Labomimus heterocerus (Coleoptera: Staphylinidae:Pselaphinae). Entomological Problems[J].2003b,33(1-2), 93-94.
Hlavac, P. & Chandler, D. S. World Catalogue of the species of Tyrini with a key to the genera (Coleoptera:Staphylinidae:Pselaphinae)[J]. Folia Heyrovskyana.2005,13(3),81-143
Hlavac, P., & Nomura, S. A Taxonomic Revision of Tyrini of the Oriental Region I. Paralasinus (Coleoptera:Staphylinidae:Pselaphinae), a new genus of Tyrina from Indochina[J]. Elytra.2001,29(1),163-174.
Hlavac, P., & Nomura, S. A taxonomic revision of Tyrini of the Oriental region III. Megatyrus (Coleoptera, Staphylinidae, Pselaphinae), a new genus of the Tyrina from China and Vietnam[J]. Elytra.2003,31(1),165-174.
Hlavac, P., Nomura, S. & Zhou, H.Z. Two new species of the genus Labomimus from China (Coleoptera:Staphylinidae, Pselaphinae) [J]. Species diversity. 2000,5 (2),149-153.
Jeannel, R. Les Pselaphides de I'Afrique orientale[J]. Memoires du Museum National d'Histoire Naturelle, (Zool.).1949,29(1),1-226.
Jeannel, R. Faune du Congo belge et du Ruanda-Urundi. II. Pselaphidae[J]. Annales du Musee du Congo Belge, (ser.8vo).1950,2 (1949),1-278.
Jeannel, R. Sur les Pselaphides (Coleoptera) de l'Inde septentrionale[J]. Bulletin of the British Museum (Natural History) (Entomology).1960,9,403-456.
Kurbatov, S.A. Le nouveau genre de Pselaphidae (Coleoptera) de l'Asie Centrale[J]. Russian Entomological Journal.1993,2 (1),51-52.
Lobl, I. Eine neue Art der Gattung Lasinus Sharp aus China (Col., Pselaphidae) [J]. Acta Rerum Naturalium Musei Nationalis Slovenici, Bratislava.1964, 10,45-48.
Lucas, R. Catalogus alphabeticus generum et subgenerum Coleopterorum orbis terrarum totius (famil., trib., subtr., sect. incl.)[J]. Pars I. Arch. Naturgesch., (A).1920,84 (1918), i-xxxi+1-696.
Motschulsky, V. de. Enumeration des nouvelles especes de Coleopteres rapportes par M. Victor Motschoulsky de son dernier voyage[J]. Bulletin de la Societe Imperiale des Naturalistes de Moscou.1851,24 (lere partie), 479-511.
Newton, A. F. & Chandler, D. S. World catalog of the genera of Pselaphidae (Coleoptera)[J]. Fieldiana, Zoology, new series.1989,53, i-iii+1-93.
Nomura, S. Taxonomical notes on Chinese species of the tyrine genus Tyrinasius (Coleoptera, Staphylinidae, Pselaphinae)[J]. Elytra.1999,27 (2), 485-498.
Panzer, G. W. F. Pselaphus mucronatus. Fauna insectorum Germaniae initia oder Deuschlands Insecten[M].1801, Heft 89, No.11.
Raffray, A. Etude sur les Pselaphides. V. Tableaux synoptiques. Notes et synonymie[J]. Revue d'Entomologie.1890a,9,81-172.
Raffray, A. Etude sur les Pselaphides. VI. Diagnoses des especes nouvelles sur lesquelles sont fondes des genres nouveaux[J]. Revue d'Entomologie[J]. 1890b,9,193-219, pls 2,3.
Raffray, A. Revision des Pselaphides des iles de Singapore et de Penang (Suite et fin) [J]. Revue d'Entomologie.1895,14,21-82, pl.2.
Raffray, A. Pselaphides nouveaux de Ceylan[J]. Annales de la Societe Entomologique de France.1901,70,27-30.
Raffray, A. Genera et catalogue des Pselaphides [J]. Annales de la Societe Entomologique de France.1904,73,1-400.
Raffray, A. Genera et catalogue des Pselaphides[J]. Annales de la Societe Entomologique de France.1905,73 (1904),401-476, pls 1-3.
Raffray, A. Especes nouvelles de Pselaphides exotiques (Coleopt.) [J]. Annales de la Societe Entomologique de France.1912,80 (1911),425-450.
Raffray, A. H. Sauter's Formosa-Ausbeute. Pselaphidae (Col.) II[J]. Supplementa Entomologica.1914,3,1-5.
Reitter, E. Versuch einer systematischen Eintheilung der Clavigeriden und Pselaphiden[J]. Verhandlungen der Naturforschenden Vereines in Brunn. 1882,20,177-211.
Schaufuss, L.W. Ueber Pselaphidengattungen[J]. Nunquam Otiosus.1877,2, 450-460.
Schmidt-Goebel, H.M. Beytrag zur einer Monographie der Pselaphen, enthaltend neue Species aus Asien[M]. Erste Lieferung. Gottlieb Haase Sohne, Prague.1838,1-16.,2 pls.
Sharp, D. Descriptions of new genera and species of Pselaphidae and Scydmaenidae from Australia and New Zealand[J]. Transactions of the Entomological Society of London.1874(4),483-518.
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