关键词: 适应, 演化, 拟态, 翅图案, 蝶类

Abstract:

Understanding the origins of animal diversity and the mechanisms of adaptive evolution is one of the central questions in modern biology. Traits such as morphology, coloration, and behavior are shaped synergistically by the developmental and genetic mechanisms and environmental factors, and the studies about their interactions have long received extensive attention. The advent of high-throughput sequencing and gene-editing technologies, particularly CRISPR-Cas9, has revolutionized evolutionary developmental biology (evo-devo) by enabling detailed functional genetic studies in non-model organisms, thus bridging the gap between genotype and phenotype. Butterflies (Lepidoptera: Papilionoidea) have become a representative model system for studying animal evolution and development, owing to their wide distribution, diverse wing patterns, and short life cycles that facilitate laboratory rearing. In recent years, extensive research on mimicry phenotypes, seasonal morphs, phenotypic plasticity, and environmental interaction has uncovered the mechanisms of adaptive evolution of complex traits. This review summarizes studies on some classical examples of adaptive evolution. Recent progress has established several research systems for resolving the genetic, developmental, and evolutionary mechanisms underlying adaptive traits: (1) For Batesian mimicry, the doublesex (dsx) gene in the genus Papilio, together with related genes, governs female-limited mimicry and plays a central role in the evolution of sexually dimorphic wing patterns. (2) Other classical systems of adaptive evolution, such as the Heliconius mimicry rings, where regulatory modifications in key genes like optix govern the convergence of wing color patterns. Comparative genomic analyses reveal that distinct Heliconius species have independently evolved similar wing phenotypes through parallel changes in the regulatory elements of the same gene, providing molecular evidence and novel insights into the mechanisms underlying convergent evolution of mimicry patterns. (3) The remarkable leaf masquerade mimicry of Kallima butterflies is governed by a genomic locus containing the cortex gene, whose regulatory architecture underlies the evolution of leaf-like wing morphology and camouflage adaptation. Together, these taxa provide comparative insights into the genetic architecture and evolutionary dynamics of anti-predation strategies. The integrative approach moves beyond correlation to causation, precisely linking genetic variation to phenotypic outcomes through functional validation. Beyond mimicry, butterflies display different seasonal morphs and phenotypic plasticity in response to environmental variation. The contrasting wing patterns of Junonia coenia and Bicyclus anynana reveal how environmental cues shape developmental trajectories, offering classical models for studying adaptive plasticity. These systems collectively illustrate the dynamic responses to nature selection. Furthermore, butterflies serve as tractable models for investigating complex organism–environment interactions, including visual perception in Papilio xuthus, host-plant coevolution in Pieris rapae, and the interplay between long-distance migration and aposematism in Danaus plexippus. The above research progress demonstrates the research trend of integrating multi-omics data (pangenome and single-cell transcriptome) and gene regulatory networks in butterfly evolutionary studies. This review also provides an outlook on butterfly evolutionary research from the perspective of technological development and expansion of research directions, thereby shedding light on new ideas for animal evolutionary research. This paradigm not only deepens our understanding of universal principles in adaptive evolution but also provides a transferable framework for conservation science, bio-inspired engineering, and physiological disease mechanisms.

Key words: Adaption, Evolutionary biology, Mimicry, Wing patterning, Butterfly