Function of flow wakes for queuing trilobites: Positioning rather than drag reduction – Criteria for drag force assessment in palaeontological CFD simulations

Trilobites are one of the most important invertebrate clades in the Palaeozoic, with significant disparity in morphology and behaviour, the latter including intriguing instances of queueing. Previous studies employed Computational Fluid Dynamics (CFD) to investigate queuing behaviour in the Devonian trilobite Trimerocephalus chopini and found drag reduction effects. Novel calculations that define a ratio between drag force and Apparent Gravity (W), along with the Submerged Froude Number (Frsub), however, reveal that the obtained drag force was practically negligible in terms of the underwater mobility of trilobites. A trilobite would start to experience difficulty in forward walking only when the relative flow speed was over 42 cm/s, which is inconsistent with the interpreted palaeoenvironment or the predicted moving speed of trilobites. Nevertheless, according to the proposed cantilever model, a trilobite had the ability to sense very minute change in fluid velocity (>7.16 μm/s). High-sensitivity mechanical sensors distributed along the body, either on the exoskeleton or limbs, empowered queuing individuals to discern the fixed self-similar pattern of the wake generated by their predecessors in the queue. In general, if a trilobite were out of the wake, the asymmetrical velocity and pressure field would aid in repositioning itself, facilitating the maintenance of migratory queues. This permitted blind trilobites to securely sense their companions, compensating for lack of long-range visual capability. This paradigm of force assessment is suitable to Computational Fluid Dynamic analyses in other extinct animal-environment interactions, offering a framework to evaluate whether drags and wakes impact more on organism's mobility (W≫0.1,Frsub≫1) or their mechanical sensors, and provides a unique cross-scale insight into animals' adaptation to palaeohydrodynamic variation.