Animals deploy diverse color-based defenses against predators, including crypsis, mimicry, aposematism, and masquerade. While crypsis, mimicry, aposematism have been extensively studied, the strategy of masquerade-where organisms imitate inedible or inanimate objects such as leaves, twigs, stones, and bird droppings-remains comparatively underexplored, particularly in adult butterflies. The Indian oakleaf butterfly ( Kallima inachus) exemplifies this phenomenon, with its wings resembling dead leaves, providing a classic example of natural selection. Although it has long been postulated that these butterflies evade predation by being misidentified as dead leaves, direct experimental evidence is lacking. In the current study, using domestic chicks as predators, we manipulated their prior experience with dead leaves (model objects) while maintaining constant exposure to butterflies to test whether dead-leaf masquerade provides a protective advantage by preventing recognition. Results showed a marked delay in the initiation of attacks by chicks familiar with dead leaves compared to those with no prior exposure or those exposed to visually altered leaves. Chicks with prior dead-leaf experience required a similar amount of time to attack the butterflies as they did to attack dead leaves. These findings provide the first empirical demonstration of dead-leaf masquerade in Kallima butterflies, shedding light on its evolutionary significance. Our study highlights the effectiveness of masquerade in inducing the misclassification of butterflies as inanimate objects, showcasing the precise mimicry achieved by these organisms when viewed in isolation from the model objects. This study advances our understanding of the evolution of masquerade and its role as a potent antipredator strategy in nature.
为了躲避捕食者的攻击,动物进化出了各种各样基于颜色的防御对策,包括隐蔽、拟态、警戒色和乔装。尽管前三者已得到了广泛研究,但乔装—即生物在其生境中将自身伪装成一种不可食或无生命的物体,如树叶、树枝、石头和鸟粪等,以误导捕食者—却被报道的相对较少,特别是对蝴蝶乔装的研究。枯叶蝶( Kallima inachus)的翅膀类似于枯叶,是自然选择的经典例子。尽管长期以来人们认为这些枯叶蝶是通过欺骗捕食者,让捕食者误认为枯叶而获得保护,且常常被认为是已确定的事实,但至今为止,还没有任何实验对其进行验证过。为了验证这一假说,我们用小鸡作为捕食者进行了捕食行为实验。通过控制小鸡对取食枯叶(模型物体)的先前经验后,再给予小鸡枯叶蝶或枯叶让其取食,以验证枯叶乔装是否可以通过干扰捕食者的识别从而为枯叶蝶提供保护的假设。结果显示,与对照组小鸡(即小鸡没有取食过任何枯叶)或取食过在视觉上改变过的枯叶的小鸡相比,有先前取食视觉上没改变过的枯叶的小鸡在攻击枯叶蝶时表现出显著的延迟反应。另外,我们还发现有先前取食枯叶经验的小鸡攻击枯叶蝶所花费的时间与它们攻击枯叶的时间无显著差异。我们的研究首次实证了枯叶蝶的枯叶乔装功能,揭示了乔装在进化中的重要意义。同时,该研究也强调了乔装在误导捕食者将枯叶蝶视为枯叶的有效性,即当蝴蝶与其模型物体被隔离观察时表现出的高度相似性。这一研究推动了我们对乔装进化的理解,并强调了其作为自然界强大的反捕食对策的作用。.
Keywords: Antipredator defense; Camouflage; Dead-leaf butterfly; Masquerade; Natural selection; Visual recognition.