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Butterflies are agile fliers which use inactive and active upstrokes (US). The active US plays a secondary role to the downstroke (DS), generating both thrust and negative vertical force. However, whether their active halfstroke function is fixed or facultative has not been clarified. We showed that during multiple backward flights of an individual, postural adjustments via body angles greater than 90°, with pitch-down and pitch-up motions in the DS and US, respectively, reoriented the stroke plane and caused the reversal of the aerodynamic functions of the halfstrokes compared with forward flight. The US and DS primarily provided weight support and horizontal force, respectively, and a leading edge vortex (LEV) was formed in both halfstrokes. The US's LEV was a Class II LEV extending from wingtip to wingtip, previously reported albeit during the DS in forward flight. The US's net force contribution increased from 32% in forward to 60% in backward flight. Likewise, US weight support increased from 8 to 85%. Despite different trajectories, body postures and force orientations among flight sequences in the global frame, the halfstroke-average forces pointed in a uniform direction relative to the body in both forward and backward flight.

The reverse flight of a monarch butterfly (Danaus plexippus) is characterized by a weight-supporting upstroke and postural changes