中国乌叶蝉亚科系统分类研究（半翅目：叶蝉科）
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摘要
CHINESE ABSTRACT
本文是关于中国乌叶蝉亚科形态、分类、系统发育和生物地理学方面的综合研究论文。内容包括乌叶蝉亚科分类的历史沿革、国内外研究概况、分类特征、系统发育、生物地理学和种类描述等方面。
形态术语基本遵从张雅林（1990）和Evans（1947）的定义。所有研究标本均为成虫针插标本，雄性外生殖器解剖采用常规方法，在Leica ZOOM2000体视解剖镜下解剖、观察与鉴定，在OLYMPUS PM-10AD显微镜下绘图，成虫外部形态特征在Nikon AFX-Ⅱ体视显微镜下完成。
乌叶蝉亚科全世界已知45属231种，世界性分布，中国已知3属34种。
本研究共整理、鉴定和描记中国乌叶蝉亚科9属81种，其中包括40新种，6新记录属，9新记录种，提出1个新异名、1个新组合。
新种：白缘宽颜乌叶蝉Ectopiocephalus whitemarginus、干刺长盾叶蝉Haranga shaftstylis、阔茎长盾叶蝉H. lata、缘脊长盾叶蝉H. ridga、柄折缘乌叶蝉Neodartus handlus、方折缘乌叶蝉N. squarus、毛折缘乌叶蝉N. piliferous、网冠新折缘叶蝉Neovulturnus reticularus、四斑新折缘叶蝉N. quadvipunctatus、端曲乌叶蝉Penthimia hamata、端弯乌叶蝉P. endcrooka、盾痕乌叶蝉P. dinta、冠沟乌叶蝉P. coronalfossa、横凸乌叶蝉P. protrdera、红褐乌叶蝉P. hyaloplasma、角凹乌叶蝉P. obtusa、角乌叶蝉P. cornicula、金顶乌叶蝉P. jindingensis、茎凸乌叶蝉P. gibberosa、蓝斑乌叶蝉P. lurida、瘤乌叶蝉P. struma、宁乌叶蝉P. ningxiaensis、膨腔乌叶蝉P. bulbosa、平背乌叶蝉P. flata、三斑乌叶蝉P. trimaculata、双斑乌叶蝉P. bimaculata、天平山乌叶蝉P. tianpingshanensis、透翅乌叶蝉P. edemata、细茎乌叶蝉P. gracilepenis、显脉乌叶蝉P. hennaveins、新月乌叶蝉P. lunata、缘齿乌叶蝉P. dentatimarginis、直干乌叶蝉P. recta、短索小乌叶蝉Tomaloides shortus、褐小乌叶蝉T. sfigurus、黑头小乌叶蝉T. blackcaputus、卵小乌叶蝉T. ovoidus、勐养小乌叶蝉T. mengyangensis、双凹小乌叶蝉T. biconcavus、眼小乌叶蝉T. bipunctatus，spp. nov.。
中国新记录属：宽颜乌叶蝉属Ectopiocephalus Kirkaldy、小乌叶蝉属Tomaloides Evans、新折缘叶蝉属Neovulturnus Kirkaldy、横脉乌叶蝉属Macutella Evans、圆头叶蝉属Chinaella Evans、长索乌叶蝉属Sidelloides Evans，n. rec.。
中国新记录种：灰褐圆头叶蝉Chinaella shephardi Evans、黑横脉乌叶蝉Macutella lutea Evans、斜冠折缘乌叶蝉Neodartus mokanshanensis Ouchi、褐斑乌叶蝉Penthimia fuscomaculosa Kwon et Lee、黑腹乌叶蝉P. mudonensis Distant、黑头乌叶蝉P.
melanocephala Motschulsky、黄斑乌叶蝉P. dorsimaculata Kwon et Lee、亮黑乌叶蝉P. distanti Baker、红额长索乌叶蝉Sidelloides histrica Evans，n. rec.。
新异名：Penthimia fulviguttata Cheng et Li 2003(=褐斑乌叶蝉Penthimia fuscomaculosa Kwon et Lee, 1978), Syn. n.。
新组合：茶新折缘叶蝉Neovulturnus testacea (Kuoh), n. comb.
系统发育运用Hennig86 (Version 1.5, Farris, 1988) 支序分析软件，选用34个性状，首次对乌叶蝉亚科中国分布的9个属及外群Magnentius共计10个属的亲缘关系进行了初步探讨，得出6个支序图与一个最简约支序图，初步建立中国分布的9个属的系统发育支序图，结果表明：1）内群是一单系群；2）乌叶蝉属是乌叶蝉亚科的原始类群；3）支序图中各相邻属间形成较好的姊妹群。
生物地理学研究根据系统发育生物地理学派的观点，对世界与中国乌叶蝉亚科起源与分布进行了简单分析与解释：1）乌叶蝉亚科起源于白垩纪早期（1.35亿年前）冈瓦纳古陆的澳大利亚板块，其它各动物区现在所呈现的格局完全是由于扩散所致；2）乌叶蝉亚科昆虫在我国主要分布于华南区和华中区；3）乌叶蝉亚科昆虫属热带、亚热带种类。
论文后部附有世界与中国乌叶蝉亚科属、种名录，支序分析图，中文索引和学名索引。
新种模式标本分别保存在西北农林科技大学昆虫博物馆（NWSUAF）、中国科学院动物研究所（IZCAS）、中山大学生命科学学院（ZSU）、南开大学生命科学学院（NKU）、中国农业大学（CAU）、中国科学院上海昆虫博物馆（SEMCAS）、天津自然历史博物馆（TJNHM）。
Systematic Study on Leafhopper Subfamily Penthimiinae
from China (Hemiptera:Cicadellidae)
The present dissertation, including introduction, history and present situation both in China and abroad, mainly deals with the morphology, phylogenetic analysis, biogeography and taxonomy on the subfamily Penthimiinae from China.
Morphological terminology was adopted principally from Zhang(1990) and Evans(1947). Male genitalia preparations were made by using 5-10% weak solution of potassium hydroxide, then washed and temporarily put in plastics plate in cavities of glycerine for observation. Dissections and identifications were performed with a Leica ZOOM2000 stereomicroscope, hand-drawing of male genitalia was carried out with OLYMPUS PM– 10AD microscope with drawing tube, and external morphological characters figured under Nikon AFX-Ⅱ stereomicroscope with drawing tube.
The subfamily Penthimiinae contains 45 genera and 231 species over the world, of which 3 genera and 34 species are known from China.
In the present paper, 9 genera and 81 species known from China are diagnosed, of which 40 species are new to science and 1 new synonym and 1 new combination are proposed, 6 genera and 9 species are recorded for the first time from China. They are listed as follows:
New species: Ectopiocephalus whitemarginus、Haranga shaftstylis、H. lata、H. ridga、Neodartus handlus、N. squarus、N. piliferous、Neovulturnus reticularus、N. quadvipunctatus、Penthimia hamata、P. endcrooka、P. dinta、P. coronalfossa、P. protrdera、P. hyaloplasma、P. obtusa、P. cornicula、P. jindingensis、P. gibberosa、P. lurida、P. struma、P. ningxiaensis、P. bulbosa、P. flata、P. trimaculata、P. bimaculata、P. tianpingshanensis、P. edemata、P. gracilepenis、P. hennaveins、P. lunata、P. dentatimarginis、P. recta、Tomaloides shortus、T. sfigurus、T. blackcaputus、T. ovoidus、T. mengyangensis、T. biconcavus、T. bipunctatus，spp. nov..
New record: Ectopiocephalus Kirkaldy, Tomaloides Evans, Neovulturnus Kirkaldy, Macutella Evans, Chinaella Evans, Sidelloides Evans, Chinaella shephardi Evans、Macutella lutea Evans、Neodartus mokanshanensis Ouchi、Penthimia fuscomaculosa Kwon et Lee、P. mudonensis Distant、P. melanocephala Motschulsky、P. dorsimaculata Kwon et Lee、P. distanti Baker、Sidelloides histrica Evans, n. rec..
New synonyms: Penthimia fulviguttata Cheng et Li 2003 ( = Penthimia fuscomaculosa Kwon et Lee, 1978), Syn. n..
New combination: Neovulturnus testacea (Kuoh), n. comb..
Phylogeny of 34 Operational Taxonomic Unit(OTU) is constructed by using Hennig86 (version 1.5), of which 9 in-group genera from China of the subfamily Penthimiinae and one outgroup genus Magnentius of the subfamily Nioniinae are included and 6 phylogenetic cladograms and 1 equivalent phylogenetic cladograms are obtained. The results show that: 1) In-group Penthimminae is a monophyletic group; 2) Penthimia was separated firstly from the cladogram and formed the most primitive group of the subfamily; 3) adjacent genera in cladograms .form sister groups.
On the basis of continental drift theory and the hypothesis of vicariance and dispersal, the biogeography analysis supports the following hypotheses: 1) The original center of the subfamily Penthimiinae was probably Laurasia in the earlier Cretaceous (135Ma) and along with the separation of this land species allocated in different areas. 2) Penthimiinae mainly distributed in the South and Centre of China which belongs to the Oriental and the Palaearctic Region; 3) the Penthimiinae is tropical and sub-tropical distribution.
All phylogenetic cladograms, checklist of species, index to Chinese names and scientific names, illustration and plates are attached at the end of the paper.
The type specimens of the new species are deposited in the following institutes: Northwest Sci-Tech University of Agriculture and Forestry (NWSUAF), Institute of Zoology, Chinese Academy of Sciences (IZCAS), Zhongshan University (ZSU), Nankai University (NKU), Chinese Agriculture University(CAU), Shanghai Entomological Museum, and Chinese Academy of
本文是关于中国乌叶蝉亚科形态、分类、系统发育和生物地理学方面的综合研究论文。内容包括乌叶蝉亚科分类的历史沿革、国内外研究概况、分类特征、系统发育、生物地理学和种类描述等方面。
形态术语基本遵从张雅林（1990）和Evans（1947）的定义。所有研究标本均为成虫针插标本，雄性外生殖器解剖采用常规方法，在Leica ZOOM2000体视解剖镜下解剖、观察与鉴定，在OLYMPUS PM-10AD显微镜下绘图，成虫外部形态特征在Nikon AFX-Ⅱ体视显微镜下完成。
乌叶蝉亚科全世界已知45属231种，世界性分布，中国已知3属34种。
本研究共整理、鉴定和描记中国乌叶蝉亚科9属81种，其中包括40新种，6新记录属，9新记录种，提出1个新异名、1个新组合。
新种：白缘宽颜乌叶蝉Ectopiocephalus whitemarginus、干刺长盾叶蝉Haranga shaftstylis、阔茎长盾叶蝉H. lata、缘脊长盾叶蝉H. ridga、柄折缘乌叶蝉Neodartus handlus、方折缘乌叶蝉N. squarus、毛折缘乌叶蝉N. piliferous、网冠新折缘叶蝉Neovulturnus reticularus、四斑新折缘叶蝉N. quadvipunctatus、端曲乌叶蝉Penthimia hamata、端弯乌叶蝉P. endcrooka、盾痕乌叶蝉P. dinta、冠沟乌叶蝉P. coronalfossa、横凸乌叶蝉P. protrdera、红褐乌叶蝉P. hyaloplasma、角凹乌叶蝉P. obtusa、角乌叶蝉P. cornicula、金顶乌叶蝉P. jindingensis、茎凸乌叶蝉P. gibberosa、蓝斑乌叶蝉P. lurida、瘤乌叶蝉P. struma、宁乌叶蝉P. ningxiaensis、膨腔乌叶蝉P. bulbosa、平背乌叶蝉P. flata、三斑乌叶蝉P. trimaculata、双斑乌叶蝉P. bimaculata、天平山乌叶蝉P. tianpingshanensis、透翅乌叶蝉P. edemata、细茎乌叶蝉P. gracilepenis、显脉乌叶蝉P. hennaveins、新月乌叶蝉P. lunata、缘齿乌叶蝉P. dentatimarginis、直干乌叶蝉P. recta、短索小乌叶蝉Tomaloides shortus、褐小乌叶蝉T. sfigurus、黑头小乌叶蝉T. blackcaputus、卵小乌叶蝉T. ovoidus、勐养小乌叶蝉T. mengyangensis、双凹小乌叶蝉T. biconcavus、眼小乌叶蝉T. bipunctatus，spp. nov.。
中国新记录属：宽颜乌叶蝉属Ectopiocephalus Kirkaldy、小乌叶蝉属Tomaloides Evans、新折缘叶蝉属Neovulturnus Kirkaldy、横脉乌叶蝉属Macutella Evans、圆头叶蝉属Chinaella Evans、长索乌叶蝉属Sidelloides Evans，n. rec.。
中国新记录种：灰褐圆头叶蝉Chinaella shephardi Evans、黑横脉乌叶蝉Macutella lutea Evans、斜冠折缘乌叶蝉Neodartus mokanshanensis Ouchi、褐斑乌叶蝉Penthimia fuscomaculosa Kwon et Lee、黑腹乌叶蝉P. mudonensis Distant、黑头乌叶蝉P.
melanocephala Motschulsky、黄斑乌叶蝉P. dorsimaculata Kwon et Lee、亮黑乌叶蝉P. distanti Baker、红额长索乌叶蝉Sidelloides histrica Evans，n. rec.。
新异名：Penthimia fulviguttata Cheng et Li 2003(=褐斑乌叶蝉Penthimia fuscomaculosa Kwon et Lee, 1978), Syn. n.。
新组合：茶新折缘叶蝉Neovulturnus testacea (Kuoh), n. comb.
系统发育运用Hennig86 (Version 1.5, Farris, 1988) 支序分析软件，选用34个性状，首次对乌叶蝉亚科中国分布的9个属及外群Magnentius共计10个属的亲缘关系进行了初步探讨，得出6个支序图与一个最简约支序图，初步建立中国分布的9个属的系统发育支序图，结果表明：1）内群是一单系群；2）乌叶蝉属是乌叶蝉亚科的原始类群；3）支序图中各相邻属间形成较好的姊妹群。
生物地理学研究根据系统发育生物地理学派的观点，对世界与中国乌叶蝉亚科起源与分布进行了简单分析与解释：1）乌叶蝉亚科起源于白垩纪早期（1.35亿年前）冈瓦纳古陆的澳大利亚板块，其它各动物区现在所呈现的格局完全是由于扩散所致；2）乌叶蝉亚科昆虫在我国主要分布于华南区和华中区；3）乌叶蝉亚科昆虫属热带、亚热带种类。
论文后部附有世界与中国乌叶蝉亚科属、种名录，支序分析图，中文索引和学名索引。
新种模式标本分别保存在西北农林科技大学昆虫博物馆（NWSUAF）、中国科学院动物研究所（IZCAS）、中山大学生命科学学院（ZSU）、南开大学生命科学学院（NKU）、中国农业大学（CAU）、中国科学院上海昆虫博物馆（SEMCAS）、天津自然历史博物馆（TJNHM）。
Systematic Study on Leafhopper Subfamily Penthimiinae
from China (Hemiptera:Cicadellidae)
The present dissertation, including introduction, history and present situation both in China and abroad, mainly deals with the morphology, phylogenetic analysis, biogeography and taxonomy on the subfamily Penthimiinae from China.
Morphological terminology was adopted principally from Zhang(1990) and Evans(1947). Male genitalia preparations were made by using 5-10% weak solution of potassium hydroxide, then washed and temporarily put in plastics plate in cavities of glycerine for observation. Dissections and identifications were performed with a Leica ZOOM2000 stereomicroscope, hand-drawing of male genitalia was carried out with OLYMPUS PM– 10AD microscope with drawing tube, and external morphological characters figured under Nikon AFX-Ⅱ stereomicroscope with drawing tube.
The subfamily Penthimiinae contains 45 genera and 231 species over the world, of which 3 genera and 34 species are known from China.
In the present paper, 9 genera and 81 species known from China are diagnosed, of which 40 species are new to science and 1 new synonym and 1 new combination are proposed, 6 genera and 9 species are recorded for the first time from China. They are listed as follows:
New species: Ectopiocephalus whitemarginus、Haranga shaftstylis、H. lata、H. ridga、Neodartus handlus、N. squarus、N. piliferous、Neovulturnus reticularus、N. quadvipunctatus、Penthimia hamata、P. endcrooka、P. dinta、P. coronalfossa、P. protrdera、P. hyaloplasma、P. obtusa、P. cornicula、P. jindingensis、P. gibberosa、P. lurida、P. struma、P. ningxiaensis、P. bulbosa、P. flata、P. trimaculata、P. bimaculata、P. tianpingshanensis、P. edemata、P. gracilepenis、P. hennaveins、P. lunata、P. dentatimarginis、P. recta、Tomaloides shortus、T. sfigurus、T. blackcaputus、T. ovoidus、T. mengyangensis、T. biconcavus、T. bipunctatus，spp. nov..
New record: Ectopiocephalus Kirkaldy, Tomaloides Evans, Neovulturnus Kirkaldy, Macutella Evans, Chinaella Evans, Sidelloides Evans, Chinaella shephardi Evans、Macutella lutea Evans、Neodartus mokanshanensis Ouchi、Penthimia fuscomaculosa Kwon et Lee、P. mudonensis Distant、P. melanocephala Motschulsky、P. dorsimaculata Kwon et Lee、P. distanti Baker、Sidelloides histrica Evans, n. rec..
New synonyms: Penthimia fulviguttata Cheng et Li 2003 ( = Penthimia fuscomaculosa Kwon et Lee, 1978), Syn. n..
New combination: Neovulturnus testacea (Kuoh), n. comb..
Phylogeny of 34 Operational Taxonomic Unit(OTU) is constructed by using Hennig86 (version 1.5), of which 9 in-group genera from China of the subfamily Penthimiinae and one outgroup genus Magnentius of the subfamily Nioniinae are included and 6 phylogenetic cladograms and 1 equivalent phylogenetic cladograms are obtained. The results show that: 1) In-group Penthimminae is a monophyletic group; 2) Penthimia was separated firstly from the cladogram and formed the most primitive group of the subfamily; 3) adjacent genera in cladograms .form sister groups.
On the basis of continental drift theory and the hypothesis of vicariance and dispersal, the biogeography analysis supports the following hypotheses: 1) The original center of the subfamily Penthimiinae was probably Laurasia in the earlier Cretaceous (135Ma) and along with the separation of this land species allocated in different areas. 2) Penthimiinae mainly distributed in the South and Centre of China which belongs to the Oriental and the Palaearctic Region; 3) the Penthimiinae is tropical and sub-tropical distribution.
All phylogenetic cladograms, checklist of species, index to Chinese names and scientific names, illustration and plates are attached at the end of the paper.
The type specimens of the new species are deposited in the following institutes: Northwest Sci-Tech University of Agriculture and Forestry (NWSUAF), Institute of Zoology, Chinese Academy of Sciences (IZCAS), Zhongshan University (ZSU), Nankai University (NKU), Chinese Agriculture University(CAU), Shanghai Entomological Museum, and Chinese Academy of
引文
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