Flies and other insects routinely land upside down on a ceiling. These inverted landing maneuvers are among the most remarkable aerobatic feats, yet the full range of these behaviors and their underlying sensorimotor processes remain largely unknown. Here, we report that successful inverted landing in flies involves a serial sequence of well-coordinated behavioral modules, consisting of an initial upward acceleration followed by rapid body rotation and leg extension, before terminating with a leg-assisted body swing pivoted around legs firmly attached to the ceiling. Statistical analyses suggest that rotational maneuvers are triggered when flies' relative retinal expansion velocity reaches a threshold. Also, flies exhibit highly variable pitch and roll rates, which are strongly correlated to and likely mediated by multiple sensory cues. When flying with higher forward or lower upward velocities, flies decrease the pitch rate but increase the degree of leg-assisted swing, thereby leveraging the transfer of body linear momentum.
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