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蚊蝎蛉科昆虫分类和系统发育分析(昆虫纲:长翅目)
Taxonomy and Phylogenetic Analysis of Bittacidae(Insecta: Mecoptera)
【作者】 陈静;
【导师】 花保祯;
【作者基本信息】 西北农林科技大学 , 农业昆虫与害虫防治, 2014, 博士
【摘要】 蚊蝎蛉科Bittacidae是长翅目Mecoptera唯一世界性分布的科,栖息于森林茂密的潮湿生境,对生态环境条件要求严格,是重要的生态指示昆虫;其幼虫具一对复眼和一个中单眼,为全变态类Holometabola昆虫所仅有,在昆虫纲Insecta系统发育中占据独特地位,是联系全变态类与半变态类Hemimetabola昆虫的重要纽带。然而,蚊蝎蛉科的分类系统迄今较为混乱,急需全面系统的分类和系统发育研究。本研究首次采用支序学方法,基于成虫44个外部形态特征,其中包括1个整体特征、7个头部特征、1个胸部特征、4个足的特征、18个翅的特征和13个雄性外生殖器特征,以小蝎蛉科Nannochoristidae的微小蝎蛉属Microchorista Byers,1974和小蝎蛉属Nannochorista Tillyard,1917为外群,借助NONA和TNT等系统发育分析软件,通过最大简约法对世界蚊蝎蛉科昆虫15属51种进行了系统发育分析。并在系统发育分析的基础上,对世界蚊蝎蛉科昆虫分类进行了全面系统的修订。系统发育分析结果表明,在蚊蝎蛉科内,双尾蚊蝎蛉属Bicaubittacus Tan&Hua,2009,曲蚊蝎蛉属Harpobittacus Gerstaecker,1885,聚蚊蝎蛉属Kalobittacus Esben-Petersen,1914,新蚊蝎蛉属Neobittacus Esben-Petersen,1914,帕兹蚊蝎蛉属Pazius Navás,1913和地蚊蝎蛉属Terrobittacus Tan&Hua,2009均为单系群,而蚊蝎蛉属Bittacus进一步被证实为复杂的并系群,与除安蚊蝎蛉属Anabittacus和孤蚊蝎蛉属Eremobittacus以外的其它属为并系关系。作为蚊蝎蛉科一个包括了许多具有不明显特征种的集合,蚊蝎蛉属显然需要进一步划分。小蚊蝎蛉属Nannobittacus Esben-Petersen,1927并系于伊森克蚊蝎蛉属Issikiella Byers,1972。Kalobittacus+(Pazius+(Nannobittacus+Issikiella)4个属单独成一分支,亲缘关系较近。安娜蚊蝎蛉属Anabittacus Kimmins,1929位于蚊蝎蛉科的最基部,表现出更多的祖征;孤蚊蝎蛉属Eremobittacus Byers,1997位于亚基部位置,仅次于Anabittacus;澳蚊蝎蛉属Austrobittacus Riek,1954,埃德瑞蚊蝎蛉属Edriobittacus Byers,1974和森蚊蝎蛉属Hylobittacus Byer,1979难以分开;金蚊蝎蛉属Orobittacus Villegas&Byers,1981与乌苏里蚊蝎蛉B. ussuriensis Plutenko,1985呈姊妹群关系。在系统发育分析结果基础上,以墨西哥的亚历山大蚊蝎蛉Bittacus alexanderi Byers,1999为模式种,建立了1新属—长肢蚊蝎蛉属Longibittacus,提出4个新组合,即亚历山大蚊蝎蛉Longibittacus alexanderi (Byers,1999) comb. nov.,班克斯蚊蝎蛉L. banksi(Esben-Petersen,1915) comb. nov.,墨西哥蚊蝎蛉L. mexicanus (Klug,1838) comb. nov.和森林蚊蝎蛉L. sylvaticus (Byers,1996) comb. nov.,均移自蚊蝎蛉属Bittacus。新蚊蝎蛉属Neobittacus曾被认为是Bittacus的异名,但系统发育分析结果表明,该属为单系群,其单系性得到较高的支持度,因此本研究支持该属的建立,建议恢复其属级单元地位。小蚊蝎蛉属Nannobittacus与伊森克蚊蝎蛉属Issikiella外形十分相似,仅翅脉有一定差异;系统发育结果表明二者构成单系群,因此,建议将二属合并,在属下建立不同的亚属或种团。金蚊蝎蛉属Orobittacus在系统发育树中的位置令人困惑,推断其原因可能是所选特征包含了较多的自衍征,而系统发育软件在分析时一般将自衍征看作无效特征,通过其它特征又难以将该属分出。因此,需要进一步寻找更多合适的特征以解决该问题。在系统发育分析的基础上,根据收集到的世界蚊蝎蛉科昆虫标本,比较全面地系统总结了世界蚊蝎蛉科昆虫的分类,修订了问题属的属征,记述了蚊蝎蛉科20属69种,并照相或绘图,编制了分属、分种检索表。另外还发现我国蚊蝎蛉5新种(其中4种已发表),分别为吊罗山蚊蝎蛉Bittacus diaoluoshanus Chen&Hua,2011,九连山蚊蝎蛉Bittacusjiulianshanus Chen et al,2013,长角蚊蝎蛉Bittacus longantennatus Chen et al,2013,具毛蚊蝎蛉Bittacus setigerus Chen et al,2013和六盘水蚊蝎蛉Bittacus liupanshuicus sp. nov.;发现2中国新记录,分别为莫纳斯特尔蚊蝎蛉Bittacus monastyrskiyi Bicha,2007rec. nov.和乌苏里蚊蝎蛉Bittacus ussuriensis Plutenko,1985rec. nov.,丰富了我国蚊蝎蛉昆虫区系。
【Abstract】 Bittacidae, a cosmopolitan family in Mecoptera, are important ecological indicators. Ingeneral, they frequent moist temperate forests with dense vegetation.Their larvae bear a pair ofcompound eyes and a median ocellus on the head, a unique character in Holometabola,implying that they occupy an important positon in the phylogeny of Insecta and a significantlinkage between the Hemimetabola and Holometabola. However, the systematics of Bittacidaehas been in chaos to date, and the taxonomic and phylogenitic researches of the familyurgently need to investigate comprehensively.The phylogenetic relationship of15genera and51species of Bittacidae was cladisticallyanalyzed using NONA and TNT for the first time, with maximum parsimony based on44external morphological characters of adults, including one character on habitus, sevencharacters on head, one character on thorax, four characters on legs,18characters on wingsand13characters in male genitalia, with Microchorista Byers,1974and NannochoristaTillyard1917in Nannochoristidae as outgroups. The taxonomy of the world Bittacidae wascomprehensively revised based on the results of the phylogenitic analyses.The phylogenetic analysis shows that the genera Bicaubittacus Tan&Hua,2009,Harpobittacus Gerstaecker,1885, Kalobittacus Esben-Petersen,1914, NeobittacusEsben-Petersen,1914, Pazius Navás,1913, and Terrobittacus Tan&Hua,2009are allmonophyletic groups. However, the genus Bittacus is reconfirmed to be a complicated gradeand is paraphyletic with the genera except Anabittacus and Eremobittacus. Bittacus needs tobe split further as a repository containing many “non-outstanding” species in Bittacidae. Thegenus Nannobittacus Esben-Petersen,1927is paraphyletic with Issikiella Byers,1972. Fourgenera, Kalobittacus+(Pazius+(Nannobittacus+Issikiella), form a clade. AnabittacusKimmins,1929is positioned in Bittacidae as the most basal lineage showing moreplesiomorphies. Eremobittacus Byers,1997is placed in the genera except for Anabittacus asthe most basal lineage. Austrobittacus Riek,1954, Edriobittacus Byers,1974, andHylobittacus Byer,1979are difficult to differentiate from each other. Orobittacus Villegas&Byers,1981is a sister group of Bittacus ussuriensis Plutenko,1985.Based on the results of the phylogenetic analysis, a new genus Longibittacus gen. nov. was erected with Bittacus alexanderi Byers,1999from Mexico as its type species.Correspondingly, four new combinations, Longibittacus alexanderi (Byers,1999) comb. nov.,L. banksi (Esben-Petersen,1915) comb. nov., L. mexicanus (Klug,1838) comb. nov. and L.sylvaticus (Byers,1996) comb. nov. were transferred from the genus Bittacus.Although the genus Neobittacus Esben-Petersen,1914was synonymized with Bittacus,the results of the phylogenetic analyses show that Neobittacus is evidently a monophyleticgroup with high branch support values. Therefore the generic status of Neobittacus isrecovered. It is suggested that the genera Nannobittacus and Issikiella need to merge into onegenus with subgenera or species groups, because they are very similar in appearence withslightly different venation, and the two genera form a paraphyletic group based on thephylogenetic analysis. The position of the genus Orobittacus in the phylogenetic tree isconfusing, which may result from the reason that the characters contain many autapmorphies,which were treated as non-informative characters by the phylogenetic softwares, and it isdifficult to seperate the genus by using non-outstanding characters. To solve this problem,more suitable characters are needed.The taxonomy of the worldwide fauna of the Bittacidae was dealt with based on theresults of phylogenetic analyses in combination with the world Bittacidae specimensexamined. Some confusing genera were revised. Sixty-nine species in twenty genera wererecorded and illustrated. Keys to genera and species were provided. Five new species (four ofwhich have been published) of Bittacus are described from China, i.e. Bittacus diaoluoshanusChen&Hua,2011; Bittacus longantennatus Chen et al.,2013; Bittacus jiulianshanus Chen etal.,2013, Bittacus setigerus Chen et al.,2013, and Bittacus liupanshuicus sp. nov. Two newdistributional records, Bittacus monastyrskiyi Bicha,2007rec. nov. and Bittacus ussuriensisPlutenko,1985rec. nov. are discovered from China for the first time.