Enemy-free space maintains swallowtail butterfly host shift

Natural enemies can be significant sources of mortality for herbivorous insects and therefore important agents of natural selection. One might expect selection to favor herbivores that escape from their natural enemies into enemy-free space. Although this is an appealing idea, it has received little empirical support, and no studies have documented enemy-free space as part of a nonagricultural, nonartificial host shift. The Alaskan swallowtail butterfly, Papilio machaon aliaska, uses as host plants a species in the family Apiaceae (Cnidium cnidiifolium) along with two Asteraceae species (Artemisia arctica and Petasites frigidus). I analyzed growth and survival of P. m. aliaska larvae in the field on the three host plants in treatments that either exposed or protected them from predators. I found that, in the presence of predators, larval survival is greater on the novel hosts (Asteraceae) than on the ancestral host (Apiaceae), but that in the absence of predators survival and growth are greater on the ancestral host. These results are a demonstration of enemy-free space as a mechanism for maintaining a naturally occurring host shift.