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Effective organismal behavior responds appropriately to changes in thesurrounding environment. Attaining this delicate balance of sensitivity andstability is a hallmark of the animal kingdom. By studying the locomotorybehavior of a simple animal (\textit{Trichoplax adhaerens}) without muscles orneurons, here, we demonstrate how monociliated epithelial cells workcollectively to give rise to an agile non-neuromuscular organism. Via directvisualization of large ciliary arrays, we report the discovery of sub-secondciliary reorientations under a rotational torque that is mediated by collectivetissue mechanics and the adhesion of cilia to the underlying substrate. In atoy model, we show a mapping of this system onto an "active-elastic resonator".This framework explains how perturbations propagate information in this arrayas linear speed traveling waves in response to mechanical stimulus. Next, weexplore the implications of parametric driving in this active-elastic resonatorand show that such driving can excite mechanical 'spikes'. These spikes incollective mode amplitudes are consistent with a system driven by parametricamplification and a saturating nonlinearity. We conduct extensive numericalexperiments to corroborate these findings within a polarized active-elasticsheet. These results indicate that periodic and stochastic forcing are valuablefor increasing the sensitivity of collective ciliary flocking. We support thesetheoretical predictions via direct experimental observation of linear speedtraveling waves which arise from the hybridization of spin and overdampeddensity waves. We map how these ciliary flocking dynamics result in agilemotility via coupling between an amplified resonator and a tuning(Goldstone-like) mode of the system. This sets the stage for how activity andelasticity can self-organize into behavior which benefits the organism as awhole.

Ciliary flocking and emergent instabilities enable collective agility in a non-neuromuscular animal