Origins and diversification of subsociality in leaf beetles (Coleoptera: Chrysomelidae: Cassidinae: Chrysomelinae)

Leaf beetles (Chrysomelidae; ~40,000 species) are commonly solitary animals but subsociality, maternal care of broods, is known in Cassidinae and Chrysomelinae. We report 11 novel records from Brazil and Peru, bringing the number of subsocial chrysomelids to 35 species in 10 genera. Two evolutionary models of chrysomelid subsociality have been proposed. One proposed three independent origins within Chrysomelinae, based on the potential phylogenetic positions of subsocial genera. The other hypothesised that an evolutionary arms race between chrysomelid prey and their predators, parasites, and parasitoids has led to an escalation of defences. Using our phylogenies, we propose that subsociality originated independently in Cassidinae and Chrysomelinae, and several times within each subfamily. Subsociality was preceded by particular behaviours. In Cassidinae, exophagous larvae with chemically offensive faecal weaponry preceded aggregated living, group defences (e.g. cycloalexy), and maternal guarding. In Chrysomelinae, offensive glandular compounds preceded ovi- and viviparity before subsociality.