Evolution of larval host plant associations and adaptive radiation in pierid butterflies

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Abstract
Butterflies in the family Pieridae (Lepidoptera: Papilionoidea) feed as larvae on plants belonging primarily to three distantly related angiosperm orders: Fabales (legumes and allied plants), Brassicales (crucifers and related plants containing mustard oil glucosides), and Santalales (‘mistletoes’). However, some utilize plants from 13 other families in a further eight orders. We investigated the evolutionary history of host plant use of the Pieridae in the context of a recent phylogenetic hypothesis of the family, using simple character optimization. Although there is a close association between host plant and butterfly higher classification, we find no evidence for cospeciation but a pattern of repeated colonization and specialization. The ancestral host of the family appears to be Fabaceae or Fabales, with multiple independent shifts to other orders, including three to Santalales. The shift to Brassicales, which contain secondary compounds (glucosinolates), promoted diversification and adaptive radiation within the subfamily Pierinae. Subsequent shifts from crucifers to mistletoes (aerial-stem hemiparasites) facilitated further diversification, and more recent shifts from mistletoes to mistletoe host trees led to exploitation of novel host plants outside the conventional three orders. Possible mechanisms underlying these host shifts are briefly discussed. In the Pierinae, a striking association between host plant, larval and adult behaviour, adult phenotype, and mimicry calls for further research into possible relationships between host specialization, plant chemistry and butterfly palatability.