Beetles, barcodes and big data: A deep dive into the phylogeny of Harpalinae (Carabidae)

Abstract The ground beetles (Carabidae) are a highly species‐rich lineage of the Coleoptera, with over half of their diversity concentrated in the ~20,000 described species in the subfamily Harpalinae sensu lato . As a presumed recent radiation lacking deeply distinct morphological divisions, their taxonomic classification has been challenging, while molecular studies remain limited in the number of genes and taxa sampled. Using ~450 mitochondrial genome sequences from across the Carabidae and the major biogeographic realms we investigate the tribal relationships in Harpalinae. Our phylogenetic analysis supports a revised system that broadly divides the harpalines into two major reciprocally monophyletic lineages, corresponding to a narrowly defined Harpalinae sensu novo and a distinct Lebiinae. Within Harpalinae, we recover well‐supported subclades that mostly represent existing tribes (e.g., Harpalini, Pterostichini, Licinini, Platynini), while clades in Lebiinae required the recognition of three new or redefined clades: Lebiini, Agrini and Odacanthini. We also establish the polyphyletic status of the ‘Truncatipennes’ defined by truncated elytra and traditionally encompassing most ‘lebiomorphs’, which are split into the Lebiinae and at least two additional lineages, corresponding to the Dryptinae and Brachininae (bombardier beetles) branching below the Harpalinae + Lebiinae clade. The mitogenome data were extended to include ~7000 species of Carabidae by adding all available cytochrome c oxidase subunit I (COI) barcodes and other legacy sequences. The resulting phylogeny broadly concurs with the tribal boundaries defined by mitogenomes and provides a curated barcode reference library for species identification. The unprecedented scale of mitogenome sequencing, combined with dense taxon sampling of barcodes, resolves a particularly complex portion of the beetle tree‐of‐life.