Time-calibrated phylogenomics reveals relationships and parallel evolution of neoteny within Elateriformia (Coleoptera) despite pervasive homoplasy

The controversy over Elateriformia relationships hampers understanding of the evolution of disparate soft-bodied and fully sclerotized phenotypes, bioluminescence, and the ontogenetic reprogramming that leads to larviform females. Elateriformia contains >50,000 species in 34 families, including fireflies, jewel, and click beetles. We sample 31 families and 195 ingroup taxa, each with up to 4,224 orthologs. Through explicit hypothesis testing, coalescent-based approaches, and evaluation of dataset properties, we resolve backbone relationships. We reconfirm the monophyly of Byrrhoidea sensu lato, rejecting the recent resurrection of Dryopoidea, and verify the reciprocally monophyletic lampyroid and elaterid clades. We characterize short internodes and gene-tree discordance, both of which are indicative of ancient rapid radiations. Counterintuitively, signals from moderately variable genes are crucial for resolving certain splits, suggesting that even subtle rate differences distinguish phylogenetically informative loci from overly conserved, less informative markers. Time-calibrated phylogenies indicate the early origin of Elateriformia (281-287 Mya), with superfamilies diverging at the Permian-Triassic boundary and diversifying throughout the Jurassic. We identify 15 independent origins of soft-bodiedness and neotenic modifications spanning the Mid Jurassic to the present. Our hypothesis provides a foundation for investigating the genetic mechanisms underlying ontogenetic reprogramming and bioluminescence.