Model tests of gliding with different hindwing configurations in the four-winged dromaeosaurid Microraptor gui

Abstract
Fossils of the remarkable dromaeosaurid Microraptor gui and relatives clearly show well-developed flight feathers on the hind limbs as well as the front limbs. No modern vertebrate has hind limbs functioning as independent, fully developed wings; so, lacking a living example, little agreement exists on the functional morphology or likely flight configuration of the hindwing. Using a detailed reconstruction based on the actual skeleton of one individual, cast in the round, we developed light-weight, three-dimensional physical models and performed glide tests with anatomically reasonable hindwing configurations. Models were tested with hindwings abducted and extended laterally, as well as with a previously described biplane configuration. Although the hip joint requires the hindwing to have at least 20 degrees of negative dihedral (anhedral), all configurations were quite stable gliders. Glide angles ranged from 3 degrees to 21 degrees with a mean estimated equilibrium angle of 13.7 degrees, giving a lift to drag ratio of 4.1:1 and a lift coefficient of 0.64. The abducted hindwing model's equilibrium glide speed corresponds to a glide speed in the living animal of 10.6 m x s(-1). Although the biplane model glided almost as well as the other models, it was structurally deficient and required an unlikely weight distribution (very heavy head) for stable gliding. Our model with laterally abducted hindwings represents a biologically and aerodynamically reasonable configuration for this four-winged gliding animal. M. gui's feathered hindwings, although effective for gliding, would have seriously hampered terrestrial locomotion.