中国弄蝶科分类与系统发育（鳞翅目：弄蝶总科）
摘要
本文对中国弄蝶科进行了分类研究,共记载4亚科82属360种,发现1中国新纪录属、2中国新纪录种,提出了3个新异名。根据标本详细记述了72属189种的外部形态特征和外生殖器特征,绘制了259张雌、雄性外生殖器图,其中53种的雌性外生殖器和1种的雄性外生殖器为首次描述;少数未见标本的属、种,根据文献对其引证和分布地进行了记载。
中国新纪录属:卡弄蝶属Carcharodus Hübner, 1819。
中国新纪录种:花卡弄蝶Carcharodus flocciferus (Zeller, 1847),库珂弄蝶Caltoris kumara (Moore, 1878)。
新异名:束带弄蝶Lobocla contractus (Leech, 1894) syn. nov. [=双带弄蝶Lobocla bifasciata (Bremer & Grey, 1853)],四纹带弄蝶Lobocla quadripunctata Fan & Wang, 2004 syn. nov. [=双带弄蝶Lobocla bifasciata (Bremer & Grey, 1853)],长刺刺胫弄蝶Baoris longistigmata Huang, 1999 syn. nov. [=刺胫弄蝶Baoris farri (Moore, 1878)]。
在分类部分的基础上,对弄蝶的69个形态特征进行了系统研究,选用了其中56个特征(包括33个外部形态特征,23个雌、雄性外生殖器特征)和3个生物学特征,对中国弄蝶科72属进行了系统发育分析,结果表明:
弄蝶科为单系群;中国弄蝶科分为4亚科,即:竖翅弄蝶亚科Coeliadinae、花弄蝶亚科Pyrginae、链弄蝶亚科Heteropterinae和弄蝶亚科Hesperiinae,它们的系统发育关系为:(Coeliadinae +(Pyrginae +(Heteropterinae + Hesperiinae))),各姐妹群关系均获得了很好的支持。
花弄蝶亚科Pyrginae可分为6个族级单元:珠弄蝶族Erynnini、花弄蝶族Pyrgini、珍弄蝶族Eudamini、彩弄蝶属群Caprona Group、星弄蝶族Celaenorrhinini和裙弄蝶族Tagiadini,各族间关系为:((Erynnini+Pyrginae)+(Eudamini+(Caprona Group+(Celaenorrhinini+Tagiadini))))。其中,珠弄蝶族Erynnini和星弄蝶族Celaenorrhinini在本次研究中得到了进一步确认;珍弄蝶族Eudamini和( Caprona Group+(Celaenorrhinini+Tagiadini)))的姐妹群关系未得到较好的支持,该族的位置需要进一步研究。
弄蝶亚科Hesperiinae可分为9个族级单元:腌翅弄蝶属群Astictopterus Group、锷弄蝶族Aeromachini、雅弄蝶属群Iambrix Group、蕉弄蝶属群Erionota Group、钩弄蝶属群Ancistroides Group、旖弄蝶族Isoteinonini、刺胫弄蝶族Baorini、黄弄蝶族Taractrocerini、弄蝶族Hesperiini,各族间关系为((Astictopterus Group+Aeromachini)+(Iambrix Group+(Erionota Group+(Ancistroides Group+(Isoteinonini+(Baorini+(Taractrocerini+Hesperiini)))))))。其中,蕉弄蝶属群Erionota Group、钩弄蝶属群Ancistroides Group、刺胫弄蝶族Baorini、黄弄蝶族Taractrocerini为单系群;旖弄蝶族Isoteinonini各属的位置及与其他属的关系未能得到很好的解决;各族级单元的姐妹群关系中,(Astictopterus Group+ Aeromachini)、(Isoteinonini+(Baorini + (Taractrocerini + Hesperiini )))、( Erionota Group+ ( Ancistroides Group+ ( Isoteinonini+ ( Baorini+(Taractrocerini + Hesperiini)))))未得到较好的支持,其余各姐妹群关系均获得了一般的支持。
同时,因缺乏标本而未纳入系统发育研究的10属,通过对相关文献中记载的特征进行分析,对它们在系统树中的位置进行了安排。
文中还编制了中国弄蝶科分亚科、族、属、种的检索表,整理提供了完整的中国弄蝶科名录。
Taxonomy of four subfamilies, 82 genera and 360 species of Hesperiidae presented in China is studied. External and genital characters of 189 species from 72 genera are described based on examinations of pinned specimens, with 259 figures of male and female genitalia illustrated, of which, one male and 53 female genitalia are described for the first time. For genera and species with material unavailable, their citation and distribution are recorded according to related literatures.
One genus and two species are discovered as new records to China, and three species are synonymized, as listed below:
New recorded genus to China: Carcharodus Hübner, 1819.
New recorded species to China: Carcharodus flocciferus (Zeller, 1847); Caltoris kumara (Moore, 1878).
New synonyms: Lobocla contractus (Leech, 1894) syn. nov. [=Lobocla bifasciata (Bremer & Grey, 1853)]; Lobocla quadripunctata Fan & Wang, 2004 syn. nov. [=Lobocla bifasciata (Bremer & Grey, 1853)]; Baoris longistigmata Huang, 1999 syn. nov. [=Baoris farri (Moore, 1878)].
Fifty six morphological characters and three biological characters are employed to analysis the phylogenetic relationship between 72 genera of Hesperiidae, using PAUP 4.0 beta 10 and Treeview 1.6.6. Monophyly of the family is strongly supported, as are the four subfamilies, with the following relationships: (Coeliadinae + (Pyrginae + (Heteropterinae + Hesperiinae))).
Six tribes of Pyrginae are recognized, of which, Erynnin and Celaenorrhinini are confirmed. Lobocla, as the only genus of Eudaminae occurring outside the Neotropical and Nearctic regions, appears as a tribe of Pyrginae in this study, but its sister relationship with (Caprona Group + (Celaenorrhinini + Tagiadini))) is poorly supported.
Nine hesperiine tribes and genus groups are partitioned with the relationship as follows: ((Astictopterus Group + Aeromachini) + (Iambrix Group + (Erionota Group + (Ancistroides Group + (Isoteinonini + (Baorini + (Taractrocerini + Hesperiini))))))). Erionota Group, Ancistroides Group, Baorini and Taractrocerini are proved to be monophyly. Sister relationships between Astictopterus Group and Aeromachini, Isoteinonini and (Baorini + (Taractrocerini + Hesperiini)), Erionota Group and (Ancistroides Group + (Isoteinonini + Baorini + (Taractrocerini + Hesperiini)))) receive week support, others have moderate support. Circumscriptions of Isoteinonini and its relationship with related genera require further elucidation.
Ten genera are not represented in the phylogenetic analysis since materials are not available, and their positions are arranged based on analyses of the characters provided in literature.
According to the results of taxonomic and phylogenetic study, keys to hesperid subfamilies, tribes, genera and species are provided. An updated checklist of Hesperiidae from China is given.
中国新纪录属:卡弄蝶属Carcharodus Hübner, 1819。
中国新纪录种:花卡弄蝶Carcharodus flocciferus (Zeller, 1847),库珂弄蝶Caltoris kumara (Moore, 1878)。
新异名:束带弄蝶Lobocla contractus (Leech, 1894) syn. nov. [=双带弄蝶Lobocla bifasciata (Bremer & Grey, 1853)],四纹带弄蝶Lobocla quadripunctata Fan & Wang, 2004 syn. nov. [=双带弄蝶Lobocla bifasciata (Bremer & Grey, 1853)],长刺刺胫弄蝶Baoris longistigmata Huang, 1999 syn. nov. [=刺胫弄蝶Baoris farri (Moore, 1878)]。
在分类部分的基础上,对弄蝶的69个形态特征进行了系统研究,选用了其中56个特征(包括33个外部形态特征,23个雌、雄性外生殖器特征)和3个生物学特征,对中国弄蝶科72属进行了系统发育分析,结果表明:
弄蝶科为单系群;中国弄蝶科分为4亚科,即:竖翅弄蝶亚科Coeliadinae、花弄蝶亚科Pyrginae、链弄蝶亚科Heteropterinae和弄蝶亚科Hesperiinae,它们的系统发育关系为:(Coeliadinae +(Pyrginae +(Heteropterinae + Hesperiinae))),各姐妹群关系均获得了很好的支持。
花弄蝶亚科Pyrginae可分为6个族级单元:珠弄蝶族Erynnini、花弄蝶族Pyrgini、珍弄蝶族Eudamini、彩弄蝶属群Caprona Group、星弄蝶族Celaenorrhinini和裙弄蝶族Tagiadini,各族间关系为:((Erynnini+Pyrginae)+(Eudamini+(Caprona Group+(Celaenorrhinini+Tagiadini))))。其中,珠弄蝶族Erynnini和星弄蝶族Celaenorrhinini在本次研究中得到了进一步确认;珍弄蝶族Eudamini和( Caprona Group+(Celaenorrhinini+Tagiadini)))的姐妹群关系未得到较好的支持,该族的位置需要进一步研究。
弄蝶亚科Hesperiinae可分为9个族级单元:腌翅弄蝶属群Astictopterus Group、锷弄蝶族Aeromachini、雅弄蝶属群Iambrix Group、蕉弄蝶属群Erionota Group、钩弄蝶属群Ancistroides Group、旖弄蝶族Isoteinonini、刺胫弄蝶族Baorini、黄弄蝶族Taractrocerini、弄蝶族Hesperiini,各族间关系为((Astictopterus Group+Aeromachini)+(Iambrix Group+(Erionota Group+(Ancistroides Group+(Isoteinonini+(Baorini+(Taractrocerini+Hesperiini)))))))。其中,蕉弄蝶属群Erionota Group、钩弄蝶属群Ancistroides Group、刺胫弄蝶族Baorini、黄弄蝶族Taractrocerini为单系群;旖弄蝶族Isoteinonini各属的位置及与其他属的关系未能得到很好的解决;各族级单元的姐妹群关系中,(Astictopterus Group+ Aeromachini)、(Isoteinonini+(Baorini + (Taractrocerini + Hesperiini )))、( Erionota Group+ ( Ancistroides Group+ ( Isoteinonini+ ( Baorini+(Taractrocerini + Hesperiini)))))未得到较好的支持,其余各姐妹群关系均获得了一般的支持。
同时,因缺乏标本而未纳入系统发育研究的10属,通过对相关文献中记载的特征进行分析,对它们在系统树中的位置进行了安排。
文中还编制了中国弄蝶科分亚科、族、属、种的检索表,整理提供了完整的中国弄蝶科名录。
Taxonomy of four subfamilies, 82 genera and 360 species of Hesperiidae presented in China is studied. External and genital characters of 189 species from 72 genera are described based on examinations of pinned specimens, with 259 figures of male and female genitalia illustrated, of which, one male and 53 female genitalia are described for the first time. For genera and species with material unavailable, their citation and distribution are recorded according to related literatures.
One genus and two species are discovered as new records to China, and three species are synonymized, as listed below:
New recorded genus to China: Carcharodus Hübner, 1819.
New recorded species to China: Carcharodus flocciferus (Zeller, 1847); Caltoris kumara (Moore, 1878).
New synonyms: Lobocla contractus (Leech, 1894) syn. nov. [=Lobocla bifasciata (Bremer & Grey, 1853)]; Lobocla quadripunctata Fan & Wang, 2004 syn. nov. [=Lobocla bifasciata (Bremer & Grey, 1853)]; Baoris longistigmata Huang, 1999 syn. nov. [=Baoris farri (Moore, 1878)].
Fifty six morphological characters and three biological characters are employed to analysis the phylogenetic relationship between 72 genera of Hesperiidae, using PAUP 4.0 beta 10 and Treeview 1.6.6. Monophyly of the family is strongly supported, as are the four subfamilies, with the following relationships: (Coeliadinae + (Pyrginae + (Heteropterinae + Hesperiinae))).
Six tribes of Pyrginae are recognized, of which, Erynnin and Celaenorrhinini are confirmed. Lobocla, as the only genus of Eudaminae occurring outside the Neotropical and Nearctic regions, appears as a tribe of Pyrginae in this study, but its sister relationship with (Caprona Group + (Celaenorrhinini + Tagiadini))) is poorly supported.
Nine hesperiine tribes and genus groups are partitioned with the relationship as follows: ((Astictopterus Group + Aeromachini) + (Iambrix Group + (Erionota Group + (Ancistroides Group + (Isoteinonini + (Baorini + (Taractrocerini + Hesperiini))))))). Erionota Group, Ancistroides Group, Baorini and Taractrocerini are proved to be monophyly. Sister relationships between Astictopterus Group and Aeromachini, Isoteinonini and (Baorini + (Taractrocerini + Hesperiini)), Erionota Group and (Ancistroides Group + (Isoteinonini + Baorini + (Taractrocerini + Hesperiini)))) receive week support, others have moderate support. Circumscriptions of Isoteinonini and its relationship with related genera require further elucidation.
Ten genera are not represented in the phylogenetic analysis since materials are not available, and their positions are arranged based on analyses of the characters provided in literature.
According to the results of taxonomic and phylogenetic study, keys to hesperid subfamilies, tribes, genera and species are provided. An updated checklist of Hesperiidae from China is given.
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