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°of negative dihedral (anhedral), all configurations were quite stable gliders. Glide angles ranged from 3°to 21°with a mean estimated equilibrium angle of 13.7°, 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·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.E vidence now exists that should settle the long-running debate over a ground-up origin of avian flight vs. the evolution of avian flight from a trees-down glider. This evidence shows that the protoavian was arboreal (1) rather than a terrestrial cursor as many have suggested (2-4). The leading protagonist in this controversy is presently a dromaeosaur, Microraptor gui, with a fully formed hind wing that is closely similar to its completely avian forewing (5), having elongate, aerodynamically advanced "primary feathers" coming off the metatarsi (5). There seems little reason to doubt the aerodynamic function of the hindwing, but there has been controversy over exactly how it was arranged and used for flight. We decided to address this problem by creating and testing models that closely replicate the anatomical features preserved in the now numerous fossils of Microraptor and its close relatives (discussed in SI Text).Primitively, early archosaurs are sprawling, with the legs set laterally and elevated at around 75°(6), a preadapted posture for gliding. Modern birds normally have the thigh elevated and sprawled to the side in different degrees; for example, it is nearly perpendicular to the midline in loons and grebes (7). This variation shows that the degree of splaying needed to use the hindlegs in gliding is not unusual when compared with that in modern birds. The absence of an antitrochanter and a supraacetabular shelf (SAC) i...