Wing-assisted incline running[edit]
The WAIR hypothesis, a version of the "cursorial model" of the evolution of avian flight, in which birds' wings originated from forelimb modifications that provided downforce, enabling the proto-birds to run up extremely steep slopes such as the trunks of trees, was prompted by observation of young
chukar chicks, and proposes that wings developed their
aerodynamic functions as a result of the need to run quickly up very steep slopes such as tree trunks, for example to escape from predators. Note that in this scenario birds need
downforce to give their feet increased grip.
[2][3] It has been argued that early birds, including
Archaeopteryx, lacked the
shoulder mechanism by which modern birds' wings produce swift, powerful upstrokes; since the downforce on which WAIR depends is generated by upstrokes, it seems that early birds were incapable of WAIR.
[4] However, a study that found lift generated from wings to be the primary factor for successfully accelerating a body toward a substrate during WAIR indicated the onset of flight ability was constrained by neuromuscular control or power output rather than by external wing morphology itself and that partially developed wings not yet capable of flight could indeed provide useful lift during WAIR.
[5] Additionally, examination of the work and power requirements for extant bird pectoralis contractile behavior during WAIR at different angles of substrate incline demonstrated incremental increases in these requirements, both as WAIR angles increased and in the transition from WAIR to flapping flight, thus providing a model for an evolutionary transition from terrestrial to aerial locomotion as transitional forms incrementally adapted to meet the work and power requirements to scale steeper and steeper inclines using WAIR and the incremental increases from WAIR to flight.
[6]
Arboreal model[edit]
This model was originally proposed in 1880 by
Othniel C. Marsh. The theory states
Archaeopteryx was a reptilian bird that soared from tree to tree. After the leap,
Archaeopteryx would then use its wings as a balancing mechanism. According to this model,
Archaeopteryx developed a gliding method to conserve energy. Even though an arboreal
Archaeopteryx exerts energy climbing the tree, an arboreal
Archaeopteryx is able to achieve higher velocities and cover greater distances during the gliding phase, which conserves more energy in the long run than a cursorial bipedal runner. Conserving energy during the gliding phase makes this a more energy-efficient model. Therefore, the benefits gained by gliding outweigh the energy used in climbing the tree. A modern behavior model to compare against would be that of the
Flying squirrel. In addition to energy conservation, arboreality is generally associated positively with survivability, at least in mammals.
[7]
The evolutionary path between arboreality and flight has been proposed through a number of hypotheses. For example, Dudley and Yanoviak proposed that animals that live in trees generally end up high enough that a fall, purposeful or otherwise, would generate enough speed for aerodynamic forces to have an affect on the body. Many animals, even those who do not fly, demonstrate the ability to right themselves and face the ground ventrally, then exhibiting behaviors that act against aerodynamic forces to slow their rate of descent in a process known as parachuting.
[7] Arboreal animals that were forced by predators or simply fell from trees that exhibited these kinds of behaviors would have been in a better position to eventually evolve capabilities that were more akin to flight as we know them today.
Researchers in support of this model have suggested that
Archaeopteryx possessed skeletal features similar to those of modern birds. The first such feature to be noted was the supposed similarity between the foot of
Archaeopteryx and that of modern perching birds. The
hallux, or modified of the first digit of the foot, was long thought to have pointed posterior to the remaining digits, as in perching birds. Therefore, researchers once concluded that
Archaeopteryx used the hallux as a balancing mechanism on tree limbs. However, study of the Thermopolis specimen of
Archeopteryx, which has the most complete foot of any known, showed that the hallux was not in fact reversed, limiting the creature's ability to perch on branches and implying a terrestrial or trunk-climbing lifestyle.
[8]
Another skeletal feature that is similar in
Archaeopteryx and modern birds is the curvature of the claws.
Archaeopteryx possessed the same claw curvature of the foot to that of perching birds. However, the claw curvature of the hand in
Archaeopteryx was similar to that in basal birds. Based upon the comparisons of modern birds to
Archaeopteryx, perching characteristics were present, signifying an arboreal habitat. The ability for takeoff and flight was originally thought to require a
supracoracoideus pulley system (SC). This system consists of a tendon joining the
humerus and
coracoid process of the
scapula allowing rotation of the humerus during the upstroke. However, this system is lacking in
Archaeopteryx. Based on experiments performed by M. Sy in 1936,
[9] it was proven that the SC pulley system was not required for flight from an elevated position but necessary for cursorial takeoff.