Temporal Precision Necessitates Wingbeat-Call Asynchrony in Actively Echolocating Bats

O_LIEcholocating bats coordinate sound production, echo reception, and flight within a closed sensorimotor loop that operates under finite propagation delays and bounded response times. Although call timing and wingbeat synchrony have been described across many behavioural contexts, it remains unclear when such coordination is temporally feasible and when it must necessarily break down. Here, I develop a constraint-based analysis that formalises how temporal feasibility limits shape behavioural coordination during active echolocation. C_LIO_LIBuilding on the responsivity framework, the model explicitly represents the ordering and timing dependencies between call emission, echo delay, sensory processing, and subsequent action, and derives conditions under which call timing can remain phase-locked to a cyclic motor rhythm such as the wingbeat. Analysis of these constraints reveals distinct coordination regimes: permissive regimes in which phase locking can emerge without dedicated coupling, and constrained regimes in which progressive phase slip or decoupling becomes unavoidable as sensory demand increases. C_LIO_LIMonte Carlo simulations show that transitions between synchrony and asynchrony arise as necessary consequences of first-principles timing constraints and bounded motor dynamics, rather than from changes in behavioural strategy. Increased motor flexibility shifts, but does not eliminate, the boundaries of synchrony-permissive regimes. Empirical observations from field studies are discussed as illustrative examples of these regimes, highlighting how apparent coupling and decoupling can both emerge from the same underlying control architecture. C_LIO_LITogether, this work identifies temporal feasibility as a governing constraint on echolocation behaviour, clarifies when buffered pseudo-closed-loop operation can masquerade as feedback control, and generates testable predictions for when and why wingbeat-call synchrony should fail in ecological contexts such as prey pursuit. More broadly, the framework situates bat echolocation within a general class of active sensing systems shaped by delayed feedback and bounded response dynamics. C_LI