Flight Mechanics and Control of Escape Manoeuvres in Hummingbirds. I. Flight Kinematics

Hummingbirds are nature’s masters of aerobatic manoeuvres. Previous research shows that hummingbirds and insects converged evolutionarily upon similar aerodynamic mechanisms and kinematics in hovering. Herein, we use three-dimensional kinematic data to begin to test for similar convergence of kinematics used for escape flight and to explore the effects of body size upon manoeuvring. We studied four hummingbird species in North America including two large species (magnificent hummingbird, Eugenes fulgens, 7.8 g, and blue-throated hummingbird, Lampornis clemenciae, 8.0 g) and two smaller species (broad-billed hummingbird, Cynanthus latirostris, 3.4 g, and black-chinned hummingbirds Archilochus alexandri, 3.1 g). Starting from a steady hover, hummingbirds consistently manoeuvred away from perceived threats using a drastic escape response that featured body pitch and roll rotations coupled with a large linear acceleration. Hummingbirds changed their flapping frequency and wing trajectory in all three degrees of freedom on a stroke-by-stroke basis, likely causing rapid and significant alteration of the magnitude and direction of aerodynamic forces. Thus it appears that the flight control of hummingbirds does not obey the ‘helicopter model’ that is valid for similar escape manoeuvres in fruit flies. Except for broad-billed hummingbirds, the hummingbirds had faster reaction times than those reported for visual feedback control in insects. The two larger hummingbird species performed pitch rotations and global-yaw turns with considerably larger magnitude than the smaller species, but roll rates and cumulative roll angles were similar among the four species

Take-off mechanics in hummingbirds (Trochilidae)

Initiating flight is challenging, and considerable effort has focused on understanding the energetics and aerodynamics of take-off for both machines and animals. For animal flight, the available evidence suggests that birds maximize their initial flight velocity using leg thrust rather than wing flapping. The smallest birds, hummingbirds (Order Apodiformes), are unique in their ability to perform sustained hovering but have proportionally small hindlimbs that could hinder generation of high leg thrust. Understanding the take-off flight of hummingbirds can provide novel insight into the take-off mechanics that will be required for micro-air vehicles. During take-off by hummingbirds, we measured hindlimb forces on a perch mounted with strain gauges and filmed wingbeat kinematics with high-speed video. Whereas other birds obtain 80–90% of their initial flight velocity using leg thrust, the leg contribution in hummingbirds was 59% during autonomous take-off. Unlike other species, hummingbirds beat their wings several times as they thrust using their hindlimbs. In a phylogenetic context, our results show that reduced body and hindlimb size in hummingbirds limits their peak acceleration during leg thrust and, ultimately, their take-off velocity. Previously, the influence of motivational state on take-off flight performance has not been investigated for any one organism. We studied the full range of motivational states by testing performance as the birds took off: (1) to initiate flight autonomously, (2) to escape a startling stimulus or (3) to aggressively chase a conspecific away from a feeder. Motivation affected performance. Escape and aggressive take-off featured decreased hindlimb contribution (46% and 47%, respectively) and increased flight velocity. When escaping, hummingbirds foreshortened their body movement prior to onset of leg thrust and began beating their wings earlier and at higher frequency. Thus, hummingbirds are capable of modulating their leg and wingbeat kinetics to increase take-off velocity

Resolution of a paradox: Hummingbird flight at high elevation does not come without a cost

Flight at high elevation is energetically demanding because of parallel reductions in air density and oxygen availability. The hovering flight of hummingbirds is one of the most energetically expensive forms of animal locomotion, but hummingbirds are nonetheless abundant at high elevations throughout the Americas. Two mechanisms enhance aerodynamic performance in high-elevation hummingbirds: increase in wing size and wing stroke amplitude during hovering. How do these changes in morphology, kinematics, and physical properties of air combine to influence the aerodynamic power requirements of flight across elevations? Here, we present data on the flight performance of 43 Andean hummingbird species as well as a 76-taxon multilocus molecular phylogeny that served as the historical framework for comparative analyses. Along a 4,000-m elevational transect, hummingbird body mass increased systematically, placing further aerodynamic demands on high-elevation taxa. However, we found that the minimum power requirements for hovering flight remain constant with respect to elevation because hummingbirds compensate sufficiently through increases in wing size and stroke amplitude. Thus, high-elevation hummingbirds are not limited in their capacity for hovering flight despite the challenges imposed by hypobaric environments. Other flight modes including vertical ascent and fast forward flight are more mechanically and energetically demanding, and we accordingly also tested for the maximum power available to hummingbirds by using a load-lifting assay. In contrast to hovering, excess power availability decreased substantially across elevations, thereby reducing the biomechanical potential for more complex flight such as competitive and escape maneuvers

Winter Conditions Influence Biological Responses of Migrating Hummingbirds

Conserving biological diversity given ongoing environmental changes requires the knowledge of how organisms respond biologically to these changes; however, we rarely have this information. This data deficiency can be addressed with coordinated monitoring programs that provide field data across temporal and spatial scales and with process-based models, which provide a method for predicting how species, in particular migrating species that face different conditions across their range, will respond to climate change. We evaluate whether environmental conditions in the wintering grounds of broad-tailed hummingbirds influence physiological and behavioral attributes of their migration. To quantify winter ground conditions, we used operative temperature as a proxy for physiological constraint, and precipitation and the normalized difference vegetation index (NDVI) as surrogates of resource availability. We measured four biological response variables: molt stage, timing of arrival at stopover sites, body mass, and fat. Consistent with our predictions, we found that birds migrating north were in earlier stages of molt and arrived at stopover sites later when NDVI was low. These results indicate that wintering conditions impact the timing and condition of birds as they migrate north. In addition, our results suggest that biologically informed environmental surrogates provide a valuable tool for predicting how climate variability across years influences the animal populations

Kinematics of hovering hummingbird flight along simulated and natural elevational gradients

Hovering flight is one of the most energetically demanding forms of animal locomotion. Despite the cost, hummingbirds regularly hover at high elevations, where flight is doubly challenging because of reduced air density and oxygen availability. We performed three laboratory experiments to examine how air density and oxygen partial pressure influence wingbeat kinematics. In the first study, we experimentally lowered air density but maintained constant oxygen partial pressure. Under these hypodense but normoxic conditions, hummingbirds increased stroke amplitude substantially and increased wingbeat frequency slightly. In the second experiment, we maintained constant air density but decreased oxygen partial pressure. Under these normodense but hypoxic conditions, hummingbirds did not alter stroke amplitude but instead reduced wingbeat frequency until they could no longer generate enough vertical force to offset body weight. In a final combined experiment, we decreased air density but increased oxygen availability, and found that the wingbeat kinematics were unaffected by supplemental oxygen. We also studied hovering and maximally loaded flight performance for 43 hummingbird species distributed along a natural elevational gradient in Peru. During free hovering flight, hummingbirds showed increased stroke amplitude interspecifically at higher elevations, mirroring the intra-individual responses in our first laboratory experiment. During loaded flight, hummingbirds increased both wingbeat frequency and wing stroke amplitude by 19% relative to free-flight values at any given elevation. We conclude that modulation of wing stroke amplitude is a major compensatory mechanism for flight in hypodense or hypobaric environments. By contrast, increases in wingbeat frequency impose substantial metabolic demands, are only elicited transiently and anaerobically, and cannot be used to generate additional sustained lift at high elevations

Of hummingbirds and helicopters: Hovering costs, competitive ability, and foraging strategies

Wing morphology and flight kinematics profoundly influence foraging costs and the overall behavioral ecology of hummingbirds. By analogy with helicopters, previous energetic studies have applied the momentum theory of aircraft propellers to estimate hovering costs from wing disc loading (WDL), a parameter incorporating wingspan (or length) and body mass. Variation in WDL has been used to elucidate differences either among hummingbird species in nectar-foraging strategies (e.g., territoriality, traplining) and dominance relations or among gender-age categories within species. We first demonstrate that WDL, as typically calculated, is an unreliable predictor of hovering (induced power) costs; predictive power is increased when calculations use wing length instead of wingspan and when actual wing stroke amplitudes are incorporated. We next evaluate the hypotheses that foraging strategy and competitive ability are functions of WDL, using our data in combination with those of published sources. Variation in hummingbird behavior cannot be easily classified using WDL and instead is correlated with a diversity of morphological and physiological traits. Evaluating selection pressures on hummingbird wings will require moving beyond wing and body mass measurements to include the assessment of the aerodynamic forces, power requirements, and power reserves of hovering, forward flight, and maneuvering. However, the WDLhelicopter dynamics model has been instrumental in calling attention to the importance of comparative wing morphology and related aerodynamics for understanding the behavioral ecology of hummingbirds

Salt intake and regulation in two passerine nectar drinkers: white-bellied sunbirds and New Holland honeyeaters

Avian nectarivores face the dilemma of having to conserve salts while consuming large volumes of a dilute, electrolyte-deficient diet. This study evaluates the responses to salt solutions and the regulation of salt intake in white-bellied sunbirds (Cinnyris talatala) and New Holland honeyeaters (Phylidonyris novaehollandiae). Birds were first offered a choice of four sucrose diets, containing no salt or 25, 50 or 75 mM NaCl. The experiment was repeated using five sucrose concentrations (0.075-0.63 M) as the base solution. Both species ingested similar amounts of all diets when fed the concentrated base solutions. However, when birds had to increase their intake to obtain enough energy on the dilute sucrose diets, there was a general avoidance of the higher salt concentrations. Through this diet switching, birds maintained constant intakes of both sucrose and sodium; the latter may contribute to absorption of their sugar diets. A second, no-choice experiment was designed to elucidate the renal concentrating abilities of these two nectarivores, by feeding them 0.63 M sucrose containing 5-200 mM NaCl over a 4-h trial. In both species, cloacal fluid osmolalities increased with diet NaCl concentration, but honeyeaters tended to retain ingested Na+, while sunbirds excreted it. Comparison of Na+ and K+ concentrations in ureteral urine and cloacal fluid showed that K+, but not Na+, was reabsorbed in the lower intestine of both species. The kidneys of sunbirds and honeyeaters, like those of hummingbirds, are well suited to diluting urine; however, they also appear to concentrate urine efficiently when necessary

Accelerational Implications of Hummingbird Display Dives

Integrative Biolog

Can scalable design of wings for flapping wing micro air vehicle be inspired by natural flyers?

Lift production is constantly a great challenge for flapping wing micro air vehicles (MAVs). Designing a workable wing, therefore, plays an essential role. Dimensional analysis is an effective and valuable tool in studying the biomechanics of flyers. In this paper, geometric similarity study is firstly presented. Then, the pw−AR ratio is defined and employed in wing performance estimation before the lumped parameter is induced and utilized in wing design. Comprehensive scaling laws on relation of wing performances for natural flyers are next investigated and developed via statistical analysis before being utilized to examine the wing design. Through geometric similarity study and statistical analysis, the results show that the aspect ratio and lumped parameter are independent on mass, and the lumped parameter is inversely proportional to the aspect ratio. The lumped parameters and aspect ratio of flapping wing MAVs correspond to the range of wing performances of natural flyers. Also, the wing performances of existing flapping wing MAVs are examined and follow the scaling laws. Last, the manufactured wings of the flapping wing MAVs are summarized. Our results will, therefore, provide a simple but powerful guideline for biologists and engineers who study the morphology of natural flyers and design flapping wing MAVs