The anatomy, taphonomy, taxonomy and systematic affinity of Markuelia: Early Cambrian to Early Ordovician scalidophorans

Markuelia is a vermiform, annulated introvertan animal known as embryonic fossils from the Lower Cambrian to Lower Ordovician. Analysis of an expanded and revised dataset for Introverta shows that the precise position of Markuelia within this clade is dependent on the taxa included. As a result, Markuelia is assigned to the scalidophoran total group to reflect uncertainty as to whether it is a stem-scalidophoran or a stem-priapulid. The taxonomy of the genus is revised to provide an improved taxonomic framework for material assigned to Markuelia. Five species are recognized: M. secunda Val’kov, M. hunanensis Dong and Donoghue, M. lauriei Haug et al., M. spinulifera sp. nov. and M. waloszeki sp. nov. Finally, the preservation of Markuelia is evaluated in the light of both the taphonomy of the fossil embryos themselves and the experimental taphonomy of the priapulid Priapulus caudatus, which has been proposed as both a close relative and an anatomical analogue of Markuelia.