Aerial righting reflexes in flightless animals

Animals that fall upside down typically engage in an aerial righting response so as to reorient dorsoventrally. This behavior can be preparatory to gliding or other controlled aerial behaviors and is ultimately necessary for a successful landing. Aerial righting reflexes have been described historically in various mammals such as cats, guinea pigs, rabbits, rats, and primates. The mechanisms whereby such righting can be accomplished depend on the size of the animal and on anatomical features associated with motion of the limbs and body. Here we apply a comparative approach to the study of aerial righting to explore the diverse strategies used for reorientation in midair. We discuss data for two species of lizards, the gecko Hemidactylus platyurus and the anole Anolis carolinensis, as well as for the first instar of the stick insect Extatosoma tiaratum, to illustrate size-dependence of this phenomenon and its relevance to subsequent aerial performance in parachuting and gliding animals. Geckos can use rotation of their large tails to reorient their bodies via conservation of angular momentum. Lizards with tails well exceeding snout-vent length, and correspondingly large tail inertia to body inertia ratios, are more effective at creating midair reorientation maneuvers. Moreover, experiments with stick insects, weighing an order of magnitude less than the lizards, suggest that aerodynamic torques acting on the limbs and body may play a dominant role in the righting process for small invertebrates. Both inertial and aerodynamic effects, therefore, can play a role in the control of aerial righting. We propose that aerial righting reflexes are widespread among arboreal vertebrates and arthropods and that they represent an important initial adaptation in the evolution of controlled aerial behavior.