Phylogenetic hypotheses of the relationships of arthropods to Precambrian and Cambrian problematic fossil taxa

A number of Vendian (latest Precambrian) body fossils have traditionally been considered arthropods or arthropodlike organisms. Several Cambrian ''weird wonders'' have also been linked with the arthropods. However, these relationships are difficult to express in traditional Linnean systematics. I present a morphological cladistic analysis of seven Vendian ''arthropodlike'' taxa compared with 21 representative Cambrian arthropods, lobopods, and weird wonders. Four arthropods from the later Phanerozoic (a pycnogonid, a monuran, and the problematic Cheloniellon and Arthropleura), five extant tardigrades, two extant kinorhynchs, and an extant priapulid, myriapod, pycnogonid, and onychophoran are also included. Monophyly of the Arthropoda is supported, but the anomalocarids and their relatives (Anomalopoda) fall out very close to the base of the traditional Arthropoda and should be included within it. The relationships among arthropods with uniramous appendages are not well resolved, but the group does not appear to be monophyletic. The biramous arthropods do form a clade and are divided into a crustaceanomorph clade and an arachnomorph clade that includes the trilobites. Most Vendian arthropodlike fossils form two clades, the Vendiamorpha and the Sprigginidae, in the arthropod stem group. The Lobopoda is a monophyletic clade with three branches: tardigrades, onychophorans, and marine lobopods. An unranked taxonomic scheme is proposed for the major clades identified here. There is no compelling reason to accept the hypothesis that the Vendian organisms included here are not metazoans.