Gnathostomulid phylogeny inferred from a combined approach of four molecular loci and morphology.

The phylogeny of the obscure metazoan phylum Gnathostomulida has previously only been addressed with cladistic analyses of morphological data. In the present study DNA sequence data from four molecular loci, including 18S rRNA, 28S rRNA, histone H3 and cytochrome c oxidase subunit I, are added to a revised morphological data matrix. The data set represents 23 gnathostomulid species that are analyzed under direct optimization using parsimony as the optimality criterion. The results obtained from analyzing the four molecular loci and combined morphological and molecular data under different parameter sets are generally very congruent, and differ only on minor points. The results clearly support gnathostomulid monophyly, as well as the basal division of Gnathostomulida into Filospermoidea and Bursovaginoidea. Filospermoidea were represented by species of Haplognathia and Cosmognathia, and generic monophyly is supported for both groups. Within Bursovaginoidea, Conophoralia (= Austrognathiidae) and Scleroperalia appear as sister groups. Monophyly of Mesognathariidae was confirmed as well, whereas the relationships between species of Gnathostomulidae and Onychognathiidae were contradicted by the molecular data when compared to morphological observations.