Hypolimnas misippus is a Batesian mimic of the toxic African Queen butterfly (Danaus chrysippus). Female H. misippus butterflies use two major wing patterning loci (M and A) to imitate three colour morphs of D. chrysippus found in different regions of Africa. In this study, we examine the evolution of the M locus and identify it as an example of adaptive atavism. This phenomenon involves a morphological reversion to an ancestral character that results in an adaptive phenotype. We show that H. misippus has re-evolved an ancestral wing pattern present in other Hypolimnas species, repurposing it for Batesian mimicry of a D. chrysippus morph. Using haplotagging, a linked-read sequencing technology, and our new analytical tool, Wrath, we discover two large transposable element (TE) insertions located at the M locus and establish that these insertions are present in the dominant allele responsible for producing mimetic phenotype. By conducting a comparative analysis involving additional Hypolimnas species, we demonstrate that the dominant allele is derived. This suggests that, in the derived allele, the TEs disrupt a cis-regulatory element, leading to the reversion to an ancestral phenotype that is then utilized for Batesian mimicry of a distinct model, a different morph of D. chrysippus. Our findings present a compelling instance of convergent evolution and adaptive atavism, in which the same pattern element has independently evolved multiple times in Hypolimnas butterflies, repeatedly playing a role in Batesian mimicry of diverse model species.

中文翻译：

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转座元件插入与泛热带蝴蝶 Hypolimnas misippus 的贝氏拟态相关

Hypolimnas misippus 是有毒的非洲皇后蝴蝶 (Danaus chrysippus) 的贝氏模仿品。雌性 H. misippus 蝴蝶使用两个主要的翅膀图案基因座（M 和 A）来模仿在非洲不同地区发现的 D. chrysippus 的三种颜色变形。在这项研究中，我们研究了 M 基因座的进化，并将其识别为适应性返祖现象的一个例子。这种现象涉及到祖先特征的形态回归，从而产生适应性表型。我们发现 H. misippus 重新进化了其他 Hypolimnas 物种中存在的祖先翅膀模式，将其重新用于 D. chrysippus 变体的贝茨拟态。使用单倍体标记（一种链接读测序技术）和我们的新分析工具 Wrath，我们发现位于 M 位点的两个大转座元件 (TE) 插入，并确定这些插入存在于负责产生模拟表型的显性等位基因中。通过对其他 Hypolimnas 物种进行比较分析，我们证明了优势等位基因的衍生。这表明，在衍生的等位基因中，TE 破坏了顺式调控元件，导致回复到祖先表型，然后将其用于独特模型（D. chrysippus 的不同形态）的贝茨拟态。我们的研究结果提供了一个令人信服的趋同进化和适应性返祖现象的实例，其中相同的模式元素在 Hypolimnas 蝴蝶中独立进化了多次，在不同模式物种的贝茨拟态中反复发挥作用。