Are chemical barriers necessary for evolution of butterfly-plant associations?

The association between heliconiine butterflies and Passion flower vines is composed of three or more subassociations, in which each Heliconius species group feeds on a different Passiflora subgenus. The relationships are consistent with the adaptive zone hypothesis of Ehrlich and Raven, which would suggest that (1) species of the subgenus Plectostemma proliferated as a result of chemical barriers to herbivory, which created a herbivore-free adaptive zone in which speciation and diversification took place, and (2) species of the H. erato-charitonia group overcame these barriers and entered a competitor-free adaptive zone, in which they proliferated and speciated with those plants as hosts. The hypothesis that plant secondary chemicals were responsible for creating such barriers to herbivory was tested using heliconiine species as bioassays, in which reduced growth rates indicated presence of chemical barriers to feeding. Contrary to expectation, plants of the subgenus Plectostemma showed little or no chemical defense against any species of heliconiine caterpillar. In contrast many plants of the "primitive" subgenus Granadilla possessed significant chemical barriers against herbivory by heliconiine larvae, excepting those species in the H. numata-melpomene species group. I concluded that chemical barriers to feeding were not responsible for proliferation and diversification in the subgenus Plectostemma, nor did chemicals create a competitor-free "adaptive zone" in which the H. erato-charitonia species-group could proliferate and speciate. Chemical barriers may have been important in the evolution of the subgenus Granadilla-heliconiine association. I suggest that plant allelochemics are only one of many possible barriers to herbivory which can help create "adaptive zones" for plants and their herbivores, and that the patterns of butterfly foodplant specialization discussed by Ehrlich and Raven (1964) are not necessarily the result of biochemical adaptation and counteradaptation.