“…Closed-loop roll control during the righting process ( Meresman et al, 2014;Yanoviak et al, 2010) combined with inertial measurements provided by the halteres (Dickinson et al, 1999;Fraenkel and Pringle, 1938). During flight, the halteres are known to oscillate up and down in antiphase with the wings (see Nalbach, 1993) and are therefore highly receptive to the state of wing activation (see Dickerson et al, 2019;Parween and Pratap, 2015;Deora et al, 2015Deora et al, , 2017Pratt et al, 2017). However, because of the extremely short duration of the righting reflex (∼49 ms) and the relatively long processing time required by the motion-processing neurons (about ∼50 ms in the fruit fly; see Warzecha and Egelhaaf, 2000;Frye, 2009), the righting reflex can be assumed to depend only on the angular body speed measurements provided by the halteres.…”