Heterochrony and the introduction of novel modes of morphogenesis during the evolution of moth choriogenesis

Choriogenesis in silkmoths (superfamily Bombycoidea) and in a sphingid moth (superfamily Sphingoidea) differ in major, but discrete, ways. In silkmoths, the predominant lamellar component assembles early in choriogenesis to form a thin framework. Subsequently, the lamellar framework is modified, first by expansion, and then by densification. Finally, ornate surface structures called aeropyle crowns form in some silkmoths, but they are absent in the species described here. In the sphingid, lamellar framework formation occurs throughout choriogenesis rather than largely during the early stages as in silkmoths. Lamellar densification occurs, but lamellar expansion and aeropyle crown formation do not. An evolutionary model is presented that accounts for the observed morphogenetic differences. Patterns of chorion protein synthesis in the sphingid differ from those in silkmoths in ways that are interpretable in light of the observed morphogenetic differences and the previously postulated functions of the proteins in silkmoths.