Locomotor mimicry in Heliconius butterflies: contrast analyses of flight morphology and kinematics

Müllerian mimicry is a mutualism involving the evolutionary convergence of colour patterns of prey on a warning signal to predators. Behavioural mimicry presumably adds complexity to the signal and makes it more difficult for Batesian mimics to parasitize it. To date, no one has quantified behavioural mimicry in Müllerian mimicry groups. However, morphological similarities among members of mimicry groups suggested that pitching oscillations of the body and wing–beat frequency (WBF) might converge with colour pattern. I compared the morphology and kinematics of four Heliconius species, which comprised two mimicry pairs. Because the mimics arose from two distinct lineages, the relative contributions of mimicry and phylogeny to variation in the species' morphologies and kinematics were examined. The positions of the centre of body mass and centre of wing mass and wing shape diverged among species within lineages, and converged among species within mimicry groups. WBF converged within mimicry groups, and it was coupled with body pitching frequency. However, body–pitching frequency was too variable to distinguish mimicry groups. Convergence in WBF may be due, at least in part, to biomechanical consequences of similarities in wing length, wing shape or the centre of wing mass among co–mimics. Nevertheless, convergence in WBF among passion–vine butterflies serves as the first evidence of behavioural mimicry in a mutualistic context.