Complex axial growth patterns in an early Cambrian trilobite from South Australia.

The exceptional fossil record of trilobites provides our best window on developmental processes in early euarthropods, but data on growth dynamics are limited. Here, we analyse post-embryonic axial growth in the Cambrian trilobite Estaingia bilobata from the Emu Bay Shale, South Australia. Using threshold models, we show that abrupt changes in growth trajectories of different body sections occurred in two phases, closely associated with the anamorphic/epimorphic and meraspid/holaspid transitions. These changes are similar to the progression to sexual maturity seen in certain extant euarthropods and suggest that the onset of maturity coincided with the commencement of the holaspid period. We also conduct hypothesis testing to reveal the likely controls of observed axial growth gradients and suggest that size may better explain growth patterns than moult stage. The two phases of allometric change in E. bilobata, as well as probable differing growth regulation in the earliest post-embryonic stages, suggest that observed body segmentation patterns in this trilobite were the result of a complex series of changing growth controls that characterized different ontogenetic intervals. This indicates that trilobite development is more complex than previously thought, even in early members of the clade.

Biomechanical analyses of Cambrian euarthropod limbs reveal their effectiveness in mastication and durophagy.

Durophagy arose in the Cambrian and greatly influenced the diversification of biomineralized defensive structures throughout the Phanerozoic. Spinose gnathobases on protopodites of Cambrian euarthropod limbs are considered key innovations for shell-crushing, yet few studies have demonstrated their effectiveness with biomechanical models. Here we present finite-element analysis models of two Cambrian trilobites with prominent gnathobases-Redlichia rex and Olenoides serratus-and compare these to the protopodites of the Cambrian euarthropod Sidneyia inexpectans and the modern American horseshoe crab, Limulus polyphemus. Results show that L. polyphemus, S. inexpectans and R. rex have broadly similar microstrain patterns, reflecting effective durophagous abilities. Conversely, low microstrain values across the O. serratus protopodite suggest that the elongate gnathobasic spines transferred minimal strain, implying that this species was less well-adapted to masticate hard prey. These results confirm that Cambrian euarthropods with transversely elongate protopodites bearing short, robust gnathobasic spines were likely durophages. Comparatively, taxa with shorter protopodites armed with long spines, such as O. serratus, were more likely restricted to a soft food diet. The prevalence of Cambrian gnathobase-bearing euarthropods and their various feeding specializations may have accelerated the development of complex trophic relationships within early animal ecosystems, especially the 'arms race' between predators and biomineralized prey.

Reassessing a cryptic history of early trilobite evolution.

Trilobites are an iconic Paleozoic group of biomineralizing marine euarthropods that appear abruptly in the fossil record (c. 521 million years ago) during the Cambrian 'explosion'. This sudden appearance has proven controversial ever since Darwin puzzled over the lack of pre-trilobitic fossils in the Origin of Species, and it has generally been assumed that trilobites must have an unobserved cryptic evolutionary history reaching back into the Precambrian. Here we review the assumptions behind this model, and suggest that a cryptic history creates significant difficulties, including the invocation of rampant convergent evolution of biomineralized structures and the abandonment of the synapomorphies uniting the clade. We show that a vicariance explanation for early Cambrian trilobite palaeobiogeographic patterns is inconsistent with factors controlling extant marine invertebrate distributions, including the increasingly-recognized importance of long-distance dispersal. We suggest that survivorship bias may explain the initial rapid diversification of trilobites, and conclude that the group's appearance at c. 521 Ma closely reflects their evolutionary origins.