Host plant defences and voltinism in European butterflies

1. With respect to seasonal availability for herbivores, plants defended by synthesising qualitative compounds differ from those protected by accumulation of quantitative macromolecules, leaf toughness, and low water and/or nutrient content. While the palatability of the former plants remains relatively constant during the season, the palatability of the latter group decreases with leaf age. 2. It was hypothesised that in seasonal temperate environments, quantitative plant defences should restrict the annual numbers of insect generations. To test this hypothesis, European butterflies were used as a model, both non-corrected regressions and tests controlling for phylogeny were carried out, and potentially confounding factors such as body size or occurrence in short-season environments were treated as covariables. 3. Non-phylogenetically controlled regressions corroborated that butterflies feeding on quantitatively protected hosts (woody plants + grasses) form fewer generations than species feeding on qualitatively protected forbs. Plant defences fitted voltinism better than butterfly size, and remained significant even after controlling for short seasons. Using independent contrasts, feeding on woody plants plus grasses, and feeding on woody plants only, predicted fewer generations. These patterns, however, applied exclusively for foliage-feeding species. 4. The association between plant defences and voltinism represents a hitherto overlooked pattern in the ecology of temperate herbivores. It may explain why large insects tend to form fewer generations and feed on structurally complex hosts, and why some species remain monovoltine although they are not restricted by short season.