Gut glands illuminate trunk segmentation in Cambrian fuxianhuiids

The euarthropod body consists of serially repeated segments with various degrees of functional specialization and regionalization [1]. Some representatives exhibit deviant metameric patterns expressed as an indirect correspondence between components of the exoskeleton, usually the number or position of dorsoventral sclerotized plates and walking legs (Supplemental Information) [1-3]. Segmental mismatch in the form of supernumerary walking legs per tergite (i.e. dorsal exoskeletal plate) is characteristic of fuxianhuiids, Cambrian euarthropods widely regarded as critical for understanding the origin of this phylum [4,5]. The broader significance of this organization remains obscure, however, due to the difficulty of distinguishing which components of the fuxianhuiid trunk reflect ancestral or derived traits. Here, we describe for the first time the presence of metameric midgut diverticulae in Fuxianhuia protensa from the Chengjiang Konservat-Lagerstätte and demonstrate that these digestive structures follow the segmentation pattern of the dorsal exoskeleton. Midgut diverticulae signal a predatory or scavenging ecology [6,7], falsifying the view of fuxianhuiids as simple mud-feeders [4]. Comparison with other euarthropods [1-3,5] indicates that fuxianhuiids possessed a unique mode of exoskeletal and visceral segmental mismatch, in which the tergites and midgut were segmentally patterned independently from the walking legs and ventral nerve cord. Our findings provide direct evidence of substantial developmental flexibility among stem-group euarthropods during the Cambrian.