Chemical defences in leaf beetles and their larvae: The ecological, evolutionary and taxonomic significance

The chemical defences of the Chrysomelinae are reviewed. Defensive glandular secretions have evolved independently in larvae and adults, and faster than the morphology of the glands. Both characters are used in a phylogenetic study of the Chrysomelini, disclosing suprageneric affinities. First, a close relationship between the Chrysomelina and <i>Phratora</i> is proposed. Secondly, <i>Leptinotarsa</i> and <i>Gonioctena</i> are probably more closely related to the Chrysolinina than to the Chrysomelina and <i>Phratora</i>. The qualitative and quantitative composition of larval and adult secretions sometimes differs between geographically isolated populations of the same species, which are therefore probably genetically distinct. The host plant shows an influence which explains discrepancies between the current classification and the chemical nature of the secretions. Some beetles sequester secondary plant metabolites or use them as precursors for their own defences. The original biosynthetic pathway has been lost in these species. Other factors which could affect the chemical diversity of the secretions are discussed: random events such as neutral mutations and genetic drift, conditioning or adaptation of predators to specific defensive mixtures, and the diversity of potential predators. The latter could help to explain the divergence in the chemical nature of the larval and adult secretions. The volatile irritants produced by the larvae are well suited to repel small arthropods, such as ants, whereas poisons associated with aposematic coloration in the adults could be directed against small vertebrates, such as birds.