Head anatomy and phylogenomics show the Carboniferous giant Arthropleura belonged to a millipede-centipede group

Abstract

The Carboniferous myriapod Arthropleura is the largest arthropod of all time, but its fossils are usually incomplete, limiting the understanding of its anatomy, ecology, and relationships. Micro–computed tomography applied to exceptionally preserved specimens from the Carboniferous Montceau-les-Mines Lagerstätte (France) reveals unprecedented details of its functional anatomy, such as the head and mouthparts. Arthropleura shares features with both millipedes and centipedes. Total-evidence phylogeny combining morphological and transcriptomic data resolves Arthropleura alone as a stem group millipede, but the inclusion of the highly incomplete Siluro-Devonian Eoarthropleura draws it deeper into the myriapod stem. Arthropleura suggests transitional morphology between clades united primarily by molecular information and underscores the value of total-evidence phylogenetics to understanding evolutionary history.