Cambrian lobopodians: A review of recent progress in our understanding of their morphology and evolution

Abstract

Lobopodians are an important group of organisms which appeared during the Cambrian Explosion. The underlying morphology is invariably a worm-like body bearing multiple pairs of legs. Yet in detail these animals preserve a range of morphologies and have attracted much paleontological attention; particularly since this assemblage probably includes the ancestors of living velvet worms (Onychophora), water bear (Tardigrada) and arthropods (Arthropoda). In recent years, knowledge of Cambrian lobopodians has increased dramatically based on numerous new records. However, there have been few comprehensive reviews of these animals since Ramsköld & Chen's study in 1998. In the present paper, new insights into Cambrian lobopodians are presented. The legs of Aysheaia pedunculata have a strong attachment with the body, like those of lobopodians in the Chengjiang Fauna. Hallucigenia fortis has a pair of eyes, two pairs of tentacles are observed in the ‘neck’ region while a bivalved head shield is unequivocally lacking. Some new characters for, and the orientation of, Hallucigenia sparsa are discussed. Longitudinal wrinkles on the body of Xenusion auerswalde are regarded here as putative muscles. Cardiodictyon sinicum bears doublure structures at the anterior margin of head and a pair of eye spots; the shape of dorsal plates is also reinterpreted. Onychodictyon has a pair of anterior appendages, but no sclerotized head shield. The affinities of Miraluolishania haikouensis are clarified and the proposal that M. haikouensis is a junior synonym of Luolishania longicruris is refuted. The large lobopodians, Kerygmachela, Jianshanopodia and Megadictyon – all with frontal appendages, gill-like limbs and tree-like or lamellate-like branches – may be swimming predators.