The color patterns on the wings of butterflies and moths are among the most complex manifestations of pattern formation in nature. The complexities of these patterns arise from the diversification of a conserved set of homologous elements known as the Nymphalid Ground Plan that can change color, shift position, expand, or disappear altogether. Recent work has shown that the anterior–posterior (AP) axis of the butterfly wing may also have an important role in the development and evolution of wing-pattern diversity. Here we characterize the AP axis by mapping expression domains of key regulatory genes onto the wing. We show that the butterfly wing can be subdivided into four primary regions, with the boundaries of these domains arising at the positions of the M1, M3, and Cu2 wing-veins. We find that the correlation among variation in the border ocelli is strongest for those within the same domain. We show how these domains may be used to determine phenotypic outcomes by surveying the frequency of color boundaries, tail development, and wing shape discontinuities across five major butterfly families: Lycaenidae, Nymphalidae, Papilionidae, Pieridae, and Riodinidae. Of the more than 200 genera we surveyed in this study, color pattern discontinuities emerge most often at the boundary veins M1, M3, and Cu2, and shape discontinuities and tails at veins M3 and Cu2. These findings reveal a hitherto unrecognized mode of evolution of patterning in the Lepidoptera.

中文翻译：

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鳞翅目翅膀的前后图案

蝴蝶和飞蛾翅膀上的颜色图案是自然界中图案形成最复杂的表现之一。这些图案的复杂性源于一组保守的同源元素的多样化，这些元素被称为蛱蝶平面图，它们可以改变颜色、移动位置、扩展或完全消失。最近的工作表明，蝴蝶翅膀的前后 (​​AP) 轴也可能在翅膀图案多样性的发展和演变中发挥重要作用。在这里，我们通过将关键调控基因的表达域映射到机翼上来表征 AP 轴。我们表明蝴蝶翅膀可以细分为四个主要区域，这些域的边界出现在 M1、M3 和 Cu2 翼静脉的位置。我们发现，对于同一域内的那些，边界 ocelli 中的变化之间的相关性最强。我们通过调查五个主要蝴蝶科的颜色边界、尾巴发育和翅膀形状不连续性的频率来展示如何使用这些域来确定表型结果：灰蝶科、蛱蝶科、蝴蝶科、粉蝶科和蝴蝶科。在我们在本研究中调查的 200 多个属中，颜色图案不连续性最常出现在边界脉 M1、M3 和 Cu2 处，形状不连续性和尾部出现在脉 M3 和 Cu2 处。这些发现揭示了鳞翅目中迄今为止未被认识的模式进化模式。五个主要蝴蝶科的尾巴发育和翅膀形状的不连续性：Lycaenidae、Nymphalidae、Papilionidae、Pieridae 和 Riodinidae。在我们在本研究中调查的 200 多个属中，颜色图案不连续性最常出现在边界脉 M1、M3 和 Cu2 处，形状不连续性和尾部出现在脉 M3 和 Cu2 处。这些发现揭示了鳞翅目中迄今为止未被认识的模式进化模式。五个主要蝴蝶科的尾巴发育和翅膀形状的不连续性：Lycaenidae、Nymphalidae、Papilionidae、Pieridae 和 Riodinidae。在我们在本研究中调查的 200 多个属中，颜色图案不连续性最常出现在边界脉 M1、M3 和 Cu2 处，形状不连续性和尾部出现在脉 M3 和 Cu2 处。这些发现揭示了鳞翅目中迄今为止未被认识的模式进化模式。