中国蚁蛉科翅几何形态学研究及其四属的分类修订
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摘要
形态学是昆虫系统分类学的基础,用先进的数学方法对形态特征进行量化描述和分析,是近年来形态学领域的发展趋势。几何形态测量学(Geometric morphometrics)的出现和发展,改变了人们对某些形态特征描述的含糊不清,准确地量化描述和分析形态特征,将大大减少对形态特征的判断误差,降低昆虫鉴定的错误率,有力地推动昆虫系统分类学的发展。
论文综述了几何形态测量学的发展,以及昆虫翅的形态学研究进展。
利用几何形态测量学方法研究了蚁蛉科翅的形态差异,通过对蚁蛉科5族22属56种476头标本翅的形态差异进行比较和分析,讨论了各族翅形态变异的主要部位和变化趋势。
通过对9种雌雄两性标本翅形态特征的比较,证实了蚁蛉科翅形态的两型现象并不普遍,对蚁蛉科大多数物种而言,两性的前后翅形态均无显著差异,因此,翅的形态数据雌雄可以通用。
通过标点-半标点法,和椭圆傅立叶法对16属28种蚁蛉的369头标本前后翅分别进行了差异显著性分析。多元方差分析(MANOVA)检测结果为,前翅：Wilk's Lambda=2.32E-08, F=52.72,p<0.01；后翅Wilk's Lambda=2.19E-07, F=59.1, p<0.01,数据表明前后翅形态特征在检测样本中差异极显著。对28个物种形成的378对组合进行Hotelling's pairwise检测,前翅检测结果,有308对物种翅形差异显著(p<0.05)；后翅的检测结果,344对物种翅形差异显著(p<0.05)。依据翅形态特征做的物种交叉验证判别分析,结果表明仅就翅轮廓而言,依据后翅的判别准确度高于依据前翅(前翅判别准确度54.5%,后翅判别准确度64.2%)。综合翅脉和翅轮廓特征分析,依据前翅的判别准确度高于依据后翅(前翅判别准确度86.2%,后翅判别准确度81.8%),这一结果表明翅脉构成的形状变异度前翅大于后翅。
本研究通过对前后翅形态特征的量化分析,探讨了雅蚁蛉属Layahima族级分类地位的争议问题,分析结果表明,雅蚁蛉属与树蚁蛉族Dendroleontini翅形态相似度高于雅蚁蛉属与囊蚁蛉族Myrmecaelurini的翅形态相似度。结论支持雅蚁蛉属Layahima隶属于树蚁蛉族Dendroleontini的观点。
本研究还对蚁蛉科的4个属进行了分类修订,通过核对模式标本,以及对文献资料的深入研究,澄清了一些此前遗留下来的问题,确定了在中国分布较广的一个物种的正确的学名是Myrmeleon trivialis Gerstaecker,1885,而不是以前多次报道用到的学名Myrmeleon formicarius Linnaeus,1767。确定了在中国被鉴定为长裳树蚁蛉的物种正确学名为长裳帛蚁蛉Bullanga florida (Navas,1913)而不是Dendroleon javanus。依据新获得的标本(包括一批模式标本)及资料,详细描述了15种蚁蛉,其中发表了2新种Dendrileon falcatus Zhan&Wang,2012, Bankisus sparsus Zhan&Wang,2012.,1中国新记录属Bankisus,确定了1新组合Hagenomyia littoralia (Miller&Stange1999) comb. nov.。提供了4属的地理分布图,并修订了各属的分种检索表。
The analysis of biological shape is a fundamental area of insect systematic research. The traditional description and measurements usually give incomplete and redundant information about the shape of a biological structure. In recent years, there is a significant change in morphological analysis with advanced quantitative measurement. This revolution was called geometric morphometic which is more effective in capturing information about the shape and result in more powerful statistical procedures for testing differences among biological shapes. The application of GM can decrease the errors in measurement and improve the development of the systematic and taxonomy
In the beginning, the development of GM and research progresses of insect wing shape were reviewed. Then the geometry of wings of Myrmleontidae including5tribes,22genera,476specimens was analyzed for discussing the variation of intra-tribes.
For testing sexual dimorphism of wing shape between male and female,9species with enough sample size of male and female were chosen for comparing wing shape. The results showed that there is no significant difference between male and female in Myrmeleontidae.
In order to investigate the taxonomic value of wing shape, forewing and hindwing from16genera,28species,369specimens of Myrmeleontidae based on landmark-semilandmark and elliptical Fourier method were analyzed. The results of MANOVA based on landmark-semilandmark method showed that the wing shapes in both forewing and hind wing had significant difference between species (FW:Wilk's Lambda=2.32E-08, F=52.72, p<0.01. HW:Wilk's Lambda=2.19E-07, F=59.1, p<0.01). Hotelling's pairwise comparision showed highly significant differences among308pairs of total378pairs and344pairs in forewing and hindwing, respectively. In cross validation test the proportion of forewings and hindwings correctly classified using elliptical Fourier method was54.5%and64.2%, respectively. It means that the outline of hindwings is more reliable than forewing in discriminant analysis. The proportion of forewings and hindwings correctly classified using landmark-semilandmark was86.2%and81.8%, respectively. It means that the analysis based on geometry of forewing is more accuracy than hindwing. The results indicated that the variation of landmarks is more than outline.
There is a controversy over the taxonomic status of genus Layahima. In order to provide more evidence to its taxonomic status, wing shapes of species from tribe Myrmecaelurini, Dendroleontini and genus Layahima were analyzed by using landmark-semilandmark method. The cluster analysis based procrustes distance showed that the genus Layahima is clearly closer to the tribe Dendroleontini rather than tribe Myrmecaelurini. The results confirm that Layahima bleongs to tribe Dendroleontini.
This research also focused on taxonomic review of four genera (Neuroptera, Myrmeleontidae) from China. Based on type specimens and original description, some taxonomic problems were solved. The identification of Myrmeleon formicarius Linnaeus,1767and Dendroleon javanus Banks,1915which distributed widely in China were therefore erroneous. Further study showed that the correct name of these specimens is Myrmeleon trivialis Gerstaecker,1885which was newly recorded from China and Bullanga florida (Navas,1913), respectively.15species from4genera were described and re-described in detail, including2new species:Dendroleon falcatus Than&Wang,2012, Bankisus sparsus Zhan&Wang,2012. One new combination was confirmed:Hagenomyia littoralia (Miller&Stange,1999) comb. nov. The distribution map and revised keys of4genera were provided.
论文综述了几何形态测量学的发展,以及昆虫翅的形态学研究进展。
利用几何形态测量学方法研究了蚁蛉科翅的形态差异,通过对蚁蛉科5族22属56种476头标本翅的形态差异进行比较和分析,讨论了各族翅形态变异的主要部位和变化趋势。
通过对9种雌雄两性标本翅形态特征的比较,证实了蚁蛉科翅形态的两型现象并不普遍,对蚁蛉科大多数物种而言,两性的前后翅形态均无显著差异,因此,翅的形态数据雌雄可以通用。
通过标点-半标点法,和椭圆傅立叶法对16属28种蚁蛉的369头标本前后翅分别进行了差异显著性分析。多元方差分析(MANOVA)检测结果为,前翅：Wilk's Lambda=2.32E-08, F=52.72,p<0.01；后翅Wilk's Lambda=2.19E-07, F=59.1, p<0.01,数据表明前后翅形态特征在检测样本中差异极显著。对28个物种形成的378对组合进行Hotelling's pairwise检测,前翅检测结果,有308对物种翅形差异显著(p<0.05)；后翅的检测结果,344对物种翅形差异显著(p<0.05)。依据翅形态特征做的物种交叉验证判别分析,结果表明仅就翅轮廓而言,依据后翅的判别准确度高于依据前翅(前翅判别准确度54.5%,后翅判别准确度64.2%)。综合翅脉和翅轮廓特征分析,依据前翅的判别准确度高于依据后翅(前翅判别准确度86.2%,后翅判别准确度81.8%),这一结果表明翅脉构成的形状变异度前翅大于后翅。
本研究通过对前后翅形态特征的量化分析,探讨了雅蚁蛉属Layahima族级分类地位的争议问题,分析结果表明,雅蚁蛉属与树蚁蛉族Dendroleontini翅形态相似度高于雅蚁蛉属与囊蚁蛉族Myrmecaelurini的翅形态相似度。结论支持雅蚁蛉属Layahima隶属于树蚁蛉族Dendroleontini的观点。
本研究还对蚁蛉科的4个属进行了分类修订,通过核对模式标本,以及对文献资料的深入研究,澄清了一些此前遗留下来的问题,确定了在中国分布较广的一个物种的正确的学名是Myrmeleon trivialis Gerstaecker,1885,而不是以前多次报道用到的学名Myrmeleon formicarius Linnaeus,1767。确定了在中国被鉴定为长裳树蚁蛉的物种正确学名为长裳帛蚁蛉Bullanga florida (Navas,1913)而不是Dendroleon javanus。依据新获得的标本(包括一批模式标本)及资料,详细描述了15种蚁蛉,其中发表了2新种Dendrileon falcatus Zhan&Wang,2012, Bankisus sparsus Zhan&Wang,2012.,1中国新记录属Bankisus,确定了1新组合Hagenomyia littoralia (Miller&Stange1999) comb. nov.。提供了4属的地理分布图,并修订了各属的分种检索表。
The analysis of biological shape is a fundamental area of insect systematic research. The traditional description and measurements usually give incomplete and redundant information about the shape of a biological structure. In recent years, there is a significant change in morphological analysis with advanced quantitative measurement. This revolution was called geometric morphometic which is more effective in capturing information about the shape and result in more powerful statistical procedures for testing differences among biological shapes. The application of GM can decrease the errors in measurement and improve the development of the systematic and taxonomy
In the beginning, the development of GM and research progresses of insect wing shape were reviewed. Then the geometry of wings of Myrmleontidae including5tribes,22genera,476specimens was analyzed for discussing the variation of intra-tribes.
For testing sexual dimorphism of wing shape between male and female,9species with enough sample size of male and female were chosen for comparing wing shape. The results showed that there is no significant difference between male and female in Myrmeleontidae.
In order to investigate the taxonomic value of wing shape, forewing and hindwing from16genera,28species,369specimens of Myrmeleontidae based on landmark-semilandmark and elliptical Fourier method were analyzed. The results of MANOVA based on landmark-semilandmark method showed that the wing shapes in both forewing and hind wing had significant difference between species (FW:Wilk's Lambda=2.32E-08, F=52.72, p<0.01. HW:Wilk's Lambda=2.19E-07, F=59.1, p<0.01). Hotelling's pairwise comparision showed highly significant differences among308pairs of total378pairs and344pairs in forewing and hindwing, respectively. In cross validation test the proportion of forewings and hindwings correctly classified using elliptical Fourier method was54.5%and64.2%, respectively. It means that the outline of hindwings is more reliable than forewing in discriminant analysis. The proportion of forewings and hindwings correctly classified using landmark-semilandmark was86.2%and81.8%, respectively. It means that the analysis based on geometry of forewing is more accuracy than hindwing. The results indicated that the variation of landmarks is more than outline.
There is a controversy over the taxonomic status of genus Layahima. In order to provide more evidence to its taxonomic status, wing shapes of species from tribe Myrmecaelurini, Dendroleontini and genus Layahima were analyzed by using landmark-semilandmark method. The cluster analysis based procrustes distance showed that the genus Layahima is clearly closer to the tribe Dendroleontini rather than tribe Myrmecaelurini. The results confirm that Layahima bleongs to tribe Dendroleontini.
This research also focused on taxonomic review of four genera (Neuroptera, Myrmeleontidae) from China. Based on type specimens and original description, some taxonomic problems were solved. The identification of Myrmeleon formicarius Linnaeus,1767and Dendroleon javanus Banks,1915which distributed widely in China were therefore erroneous. Further study showed that the correct name of these specimens is Myrmeleon trivialis Gerstaecker,1885which was newly recorded from China and Bullanga florida (Navas,1913), respectively.15species from4genera were described and re-described in detail, including2new species:Dendroleon falcatus Than&Wang,2012, Bankisus sparsus Zhan&Wang,2012. One new combination was confirmed:Hagenomyia littoralia (Miller&Stange,1999) comb. nov. The distribution map and revised keys of4genera were provided.
引文
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