Allometry of maximum vertical force production during hovering flight of neotropical orchid bees (Apidae: Euglossini)

Abstract
SUMMARY The ability of orchid bees to generate vertical forces was evaluated using a load-lifting method that imposed asymptotically increasing loads during ascending flight, ultimately eliciting maximum forces while hovering. Among 11 orchid bee species varying by approximately an order of magnitude in body mass, the capacity to produce vertical forces expressed relative either to body weight or to flight muscle weight declined linearly with increased body mass. Allometric analysis of log-transformed data, by contrast, found maximum vertical force to scale isometrically with body mass, but also to exhibit a slightly negative allometry with respect to flight muscle mass. Maximum stroke amplitude at limiting loads averaged 140° and was remarkably constant among species, a result consistent with anatomical constraints of the hymenopteran thorax on wing motions. By contrast, wing-beat frequencies during maximum performance declined with increasing body mass. Repeated lifting by individual bees reduced performance only when the number of consecutive lifts exceeded 15. Variation in linear mass density of the lifted load did not systematically alter performance estimates, although measurements on one species in two consecutive years at different thermal environments yielded significant differences in estimates of maximum force production. These findings suggest an adverse scaling of vertical force production at greater body mass even if flight muscle mass scales isometrically.