The physiology and biomechanics of avian flight at high altitude

Many birds fly at high altitude, either during long-distance flights or by virtue of residence in high-elevation habitats. Among the many environmental features that vary systematically with altitude, five have significant consequences for avian flight performance: ambient wind speeds, air temperature, humidity, oxygen availability, and air density. During migratory flights, birds select flight altitudes that minimize energy expenditure via selection of advantageous tail- and cross-winds. Oxygen partial pressure decreases substantially to as little as 26% of sea-level values for the highest altitudes at which birds migrate, whereas many taxa reside above 3000 meters in hypoxic air. Birds exhibit numerous adaptations in pulmonary, cardiovascular, and muscular systems to alleviate such hypoxia. The systematic decrease in air density with altitude can lead to a benefit for forward flight through reduced drag but imposes an increased aerodynamic demand for hovering by degrading lift production and simultaneously elevating the induced power requirements of flight. This effect has been well-studied in the hovering flight of hummingbirds, which occur throughout high-elevation habitats in the western hemisphere. Phylogenetically controlled studies have shown that hummingbirds compensate morphologically for such hypodense air through relative increases in wing size, and kinematically via increased stroke amplitude during the wingbeat. Such compensatory mechanisms result in fairly constant power requirements for hovering at different elevations, but decrease the margin of excess power available for other flight behaviors.