Integrating morphology and kinematics in the scaling of hummingbird hovering metabolic rate and efficiency.

Wing kinematics and morphology are influential upon the aerodynamics of flight. However, there is a lack of studies linking these variables to metabolic costs, particularly in the context of morphological adaptation to body size. Furthermore, the conversion efficiency from chemical energy into movement by the muscles (mechanochemical efficiency) scales with mass in terrestrial quadrupeds, but this scaling relationship has not been demonstrated within flying vertebrates. Positive scaling of efficiency with body size may reduce the metabolic costs of flight for relatively larger species. Here, we assembled a dataset of morphological, kinematic, and metabolic data on hovering hummingbirds to explore the influence of wing morphology, efficiency, and mass on hovering metabolic rate (HMR). We hypothesize that HMR would decline with increasing wing size, after accounting for mass. Furthermore, we hypothesize that efficiency will increase with mass, similarly to other forms of locomotion. We do not find a relationship between relative wing size and HMR, and instead find that the cost of each wingbeat increases hyperallometrically while wingbeat frequency declines with increasing mass. This suggests that increasing wing size is metabolically favourable over cycle frequency with increasing mass. Further benefits are offered to larger hummingbirds owing to the positive scaling of efficiency.

Wingbeat kinematics and energetics during weightlifting in hovering hummingbirds across an elevational gradient.

Hummingbirds differentially modify flight kinematics in response to the type of challenge imposed. Weightlifting is associated with increases in stroke amplitude (the angle swept by the wings) to increase the angular velocity of the wings and generate the requisite lift, but only up to 160°. Conversely, flight in hypodense air is accomplished by increasing the angular velocity of the wing through increases in wingbeat frequency and stroke amplitudes, with larger increases in amplitude than seen in weightlifting flight. The kinematic differences between these two challenges may be facilitated by the lower energetic costs associated with overcoming drag and inertial forces over the wing during hypodense flight. Thus, we hypothesized that energetic expenditure is what limits the kinematics of weightlifting flight, with lower air densities permitting increases in angular velocity at comparatively lower costs. To explore the kinematic and energetic effects of air density and weightlifting on hovering flight performance, video and respirometric recordings of weightlifting were performed on four species of hummingbirds across an elevational gradient. Contrary to our hypothesis, wingbeat frequency did not vary due to elevation. Instead, wingbeat frequency seems to increase depending on the power requirements for sustaining hovering flight. Furthermore, metabolic rates during hovering increased with angular velocity alone, independent of elevation. Thus, it appears that the differential responses to flight challenges are not driven by variation in the flight media.