Protaspis larva of an aglaspidid-like arthropod from the Ordovician of Siberia and its habitat.

A fossil larva lacking segmentation of the calcified carapace, closely resembling the trilobite protaspis, has been found associated with other skeletal elements of an angarocaridid Girardevia species in the mid Darriwilian of central Siberia. The presence of protaspis larvae in the angarocaridids, generally believed to represent a branch of the Aglaspidida, supports their proximity to trilobites and proves a low position on the arthropod phylogenetic tree but does not necessarily contradict the chelicerate affinity. The cephalic appendages of angarocaridids bore massive gnathobases with detachable spines, closely similar to those known in extant xiphosurans and in their probable Cambrian relatives. The stratigraphic succession of the angarocaridids, their phosphatized cuticle pieces being abundant in the Ordovician strata of Siberia, shows a gradual improvement of mechanical resistance of their carapaces, eventually resulting in a honeycomb structure. The associated benthic mollusc assemblage is dominated with the bellerophontids showing high mortality at metamorphosis and only the limpet-like Pterotheca, infaunal bivalves, and scaphopods being able to survive this in a substantial number. This suggests a strong selective pressure from predators equipped with well-skeletonised oral apparatuses able to crush mineralized body covers of their prey. Possibly, these were some of the associated conodonts of appropriate size and co-evolving towards their ability to crush more and more resistant cuticle. Less likely candidates for durophagy are endoceratid or orthoceratid cephalopods. Also the angarocaridids themselves, equipped with robust gnathobases of cephalic appendages, apparently predated on benthic shelly animals.