Laboratory of Genetics, Department of Biology, University of Turku, FIN-20014 Turku, Finland.
Mol Phylogenet Evol. 2010 Jun;55(3):929-38. doi: 10.1016/j.ympev.2010.01.025. Epub 2010 Feb 1.
In order to facilitate the study of the evolution of female flightlessness among the geometrid subfamily Ennominae (Lepidoptera, Geometridae), we carried out a phylogenetic analysis based on a morphological data matrix, and DNA sequences. We used seven nuclear gene fragments, elongation factor 1alpha (EF-1alpha), wingless (wgl), isocitrate dehydrogenase (IDH), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), ribosomal protein S5 (RpS5) and segments D1 and D2 of the 28S rRNA gene, and one mitochondrial gene fragment, cytochrome oxidase subunit I (COI). Sampling included 55 species of Ennominae covering all tribes with flightless females of the Holarctic boreal zone, and some other geometrids used as outgroups. Our results clearly confirmed that Ennominae (including Alsophila of the traditional subfamily Alsophilinae) is a monophyletic group, as well as supported the previously established morphology-based division of Ennominae into "ennomine" and "boarmiine" groups of genera. A number of taxonomic ambiguities were resolved but the monophyly of the traditionally recognised tribe Bistonini, comprising a number of flightless species, remained ambiguous. Bistonini is thus suggested to be subsumed to the tribe Boarmiini in the broad sense. Indeed, an analysis of timing of divergence suggested that Boarmiini s. lat. rapidly diversified in the late Oligocene/early Miocene. Within the Ennominae, seven independent origins of female flightlessness were revealed facilitating phylogenetic comparative analyses to be performed in search of causes and consequences of this phenomenon. The present phylogenetic hypothesis supports the conclusions of the "adaptive story", a hypothesis of the sequence of evolutionary events leading to flightlessness, we have presented earlier (Snäll et al., 2007). In particular, in the "boarmiine" group, the tribe Boarmiini s. lat. clearly represents a group of geometrids in which female flightlessness has evolved more frequently than in any other tribes, suggesting that this clade has likely been predisposed to evolutionary events leading to the manifestation of female flightlessness. The ancestor of the wing-reduced Ennominae has likely been a winged but slow flying forest moth feeding polyphagously on deciduous trees.
为了方便研究 Ennominae 亚科（鳞翅目，尺蛾科）中雌性无翅的进化，我们基于形态学数据矩阵和 DNA 序列进行了系统发育分析。我们使用了七个核基因片段，包括延伸因子 1α（EF-1α）、无翅基因（wgl）、异柠檬酸脱氢酶（IDH）、甘油醛-3-磷酸脱氢酶（GAPDH）、核糖体蛋白 S5（RpS5）以及 28S rRNA 基因的 D1 和 D2 片段，以及一个线粒体基因片段，细胞色素氧化酶亚基 I（COI）。采样包括了 55 种 Ennominae 物种，涵盖了全北极地区有雌性无翅的所有部落，以及一些其他尺蛾科作为外群。我们的结果清楚地证实，Ennominae（包括传统亚科 Alsophilinae 的 Alsophila）是一个单系群，并且支持了以前基于形态学的 Ennominae 分类，将其分为“ennomine”和“boarmiine”两个属群。一些分类学上的歧义得到了解决，但传统上被认为是有雌性无翅的 Bistonini 部落的单系性仍然存在疑问。因此，Bistonini 被建议归入广义的 Boarmiini 部落。事实上，分化时间的分析表明，Boarmiini s. lat. 在晚渐新世/早中新世迅速多样化。在 Ennominae 内部，揭示了雌性无翅的七个独立起源，这使得能够进行系统发育比较分析，以寻找这种现象的原因和后果。目前的系统发育假说支持我们早些时候提出的“适应性故事”的结论，这是一个导致无翅的进化事件序列的假说（Snäll 等人，2007）。特别是，在“boarmiine”组中，广义的 Boarmiini 部落显然代表了一个几何形物种群体，其中雌性无翅的进化比任何其他部落都更为频繁，这表明这个分支很可能已经倾向于导致雌性无翅表现的进化事件。翅膀退化的 Ennominae 的祖先很可能是一种有翅膀但飞行缓慢的森林飞蛾，以落叶树为食，取食多种食物。
