The morphology and phylogenetic position of the Cambrian lobopodian Diania cactiformis

Cambrian lobopodians have attracted much attention in that they are considered to have close affinities with the origin of arthropods, and recent studies agree that they sample the stem-groups of both Arthropoda and Onychophora. A new lobopodian species, Diania cactiformis Liu et al., 2011, was recently reported from the Chengjiang Lagerstätte (south-west China; Cambrian Series 2). This animal was interpreted as bearing arthropod-like articulated appendages and was resolved as the sister taxon to Arthropoda in a cladistic analysis. A reanalysis of the published dataset did not reproduce the phylogenetic placement of D. cactiformis, and the interpretation of its appendage morphology is controversial. New observations on D. cactiformis are made using newly collected material, rejecting the evidence for sclerotized, segmented and articulated appendages; instead the appendages compare more closely to lobopods. The criteria for defining arthropodized appendages are further discussed through comparative studies among fossil lobopodians, anomalocaridids and arthropods. Our study challenges the previous description of anteroposterior orientation, terminal claws and trunk tagmosis as expressed by differentiated anterior and posterior appendages. A synthesis of four recent datasets for lobopodians and stem-group arthropods is used to revisit the phylogeny of stem arthropods. Irrespective of how characters are weighted, D. cactiformis is resolved at a much more basal position to arthropods, indicating that the morphology of this animal's appendages does not fundamentally inform on the evolution of arthropodization.