Animals move in ‘modes’ where movement patterns relate to specific behaviours. Despite much work on the movement of butterflies, their behavioural modes are relatively unexplored. Here we analysed the behaviour of the model butterfly species the meadow brown, Maniola jurtina. We identified modes in both sexes and across habitats varying in resource density. We found that, in nectar-rich habitats, males had more diverse behaviour than females, engaging in a unique ‘high-flight’ mode associated with mate search, whereas females were primarily nectaring or inactive. In nectar-poor habitats, both sexes were similar, switching between flight and inactivity. We also identified the movement parameters of the modes, finding that, for both sexes, movements associated with nectaring were slower and more tortuous and, for males, the mode associated with mate searching was straighter and faster. Using an individual-based random-walk model, we investigated the effects of behaviour on movement predictions by comparing a mode-switching model with a version including intraspecific variation and another assuming homogeneity between individuals. For both sexes, including modes affected the mean and shape of the displacement rate compared to models assuming homogeneity, although for females modes increased displacement 1.5 times while for males they decreased it by a third. Both models also differed substantially from models assuming intraspecific variation. Finally, using a new model of search behaviour we investigated the general conditions under which individuals should engage in an exclusive search for host plants or receptive females. Parameterized for M. jurtina, the model predicted males should engage exclusively in mate search, but females only when searching is very efficient. The model provides a framework for analysing the searching behaviour of other butterfly species.

中文翻译：

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蝴蝶的行为模式：它们对运动和搜索行为的影响

动物以“模式”移动，其中移动模式与特定行为相关。尽管对蝴蝶的运动有很多研究，但它们的行为模式相对来说还没有被探索。在这里，我们分析了模式蝴蝶物种草甸棕色 Maniola jurtina 的行为。我们确定了两种性别和资源密度不同的栖息地的模式。我们发现，在富含花蜜的栖息地，雄性比雌性的行为更加多样化，参与与配偶搜索相关的独特“高飞”模式，而雌性主要是在采蜜或不活动。在缺乏花蜜的栖息地，两性相似，在逃跑和不活动之间切换。我们还确定了这些模式的运动参数，发现对于两性来说，与泌乳相关的运动更慢、更曲折，而对于雄性来说，与配偶搜索相关的模式更直接、更快。使用基于个体的随机游走模型，我们通过将模式切换模型与包括种内变异和另一个假设个体之间的同质性的版本进行比较，研究了行为对运动预测的影响。对于两性，与假设同质的模型相比，包括模式影响了位移率的平均值和形状，尽管对于女性，模式增加了 1.5 倍的位移，而对于男性，它们减少了三分之一。两种模型也与假设种内变异的模型有很大不同。最后，使用一种新的搜索行为模型，我们调查了个人应该专门搜索寄主植物或接受雌性的一般条件。参数化 M. jurtina，该模型预测男性应该专门从事配偶搜索，但女性仅在搜索非常有效时才参与。该模型为分析其他蝴蝶物种的搜索行为提供了一个框架。