Anna's Hummingbirds Research Articles

Ascending flight and decelerating vertical glides in Anna's hummingbirds.

Hummingbirds are observationally well known for their capacity to vertically ascend whilst hovering, but the underlying mechanics and possible energetic limits to ascent rates are unclear. Decelerations during vertical ascent to a fixed target may also be associated with specific visual responses to regulate the body's trajectory. Here, we studied climbing flight and subsequent deceleration in male Anna's hummingbirds (Calypte anna) over an approximately 2m vertical distance. Birds reached vertical speeds and accelerations up to ∼4 ms-1 and 10 m s-2, respectively, through the use of flapping frequencies as high as 56 Hz and stroke amplitudes slightly greater than 180deg. Total mass-specific power at maximal ascent speed was up to 92Wkg-1 body mass. Near the end of the ascending trajectory, all individuals decelerated ballistically via cessation of flapping and folding of wings over the body without losing control, a behavior termed here a vertical glide. Visual modulation of the deceleration trajectory during ascent was indicated by a constant value (∼0.75) for the first derivative of the time-to-contact to target. Our results indicate that hummingbirds in rapid vertical ascent expended near-maximal power output during flight, but also tightly controlled their subsequent deceleration during the vertical glide.

Trace element contamination in feather and tissue samples from Anna’s hummingbirds

Trace element contamination (17 elements; Be, V, Cr, Mn, Fe, Co, Ni, Cu, Zn, As, Se, Mo, Cd, Ba, Hg, Tl, and Pb) of live (feather samples only) and deceased (feather and tissue samples) Anna's hummingbirds (Calypte anna) was evaluated. Samples were analyzed using inductively coupled plasma-mass spectrometry (ICP-MS; 17 elements) and atomic absorption spectrophotometry (Hg only). Mean plus one standard deviation (SD) was considered the benchmark, and concentrations above the mean+1 SD were considered elevated above normal. Contour feathers were sampled from live birds of varying age, sex, and California locations. In order to reduce thermal impacts, minimal feathers were taken from live birds, therefore a novel method was developed for preparation of low mass feather samples for ICP-MS analysis. The study found that the novel feather preparation method enabled small mass feather samples to be analyzed for trace elements using ICP-MS. For feather samples from live birds, all trace elements, with the exception of beryllium, had concentrations above the mean+1 SD. Important risk factors for elevated trace element concentrations in feathers of live birds were age for iron, zinc, and arsenic, and location for iron, manganese, zinc, and selenium. For samples from deceased birds, ICP-MS results from body and tail feathers were correlated for Fe, Zn, and Pb, and feather concentrations were correlated with renal (Fe, Zn, Pb) or hepatic (Hg) tissue concentrations. Results for AA spectrophotometry analyzed samples from deceased birds further supported the ICP-MS findings where a strong correlation between mercury concentrations in feather and tissue (pectoral muscle) samples was found. These study results support that sampling feathers from live free-ranging hummingbirds might be a useful, non-lethal sampling method for evaluating trace element exposure and provides a sampling alternative since their small body size limits traditional sampling of blood and tissues. The results from this study provide a benchmark for the distribution of trace element concentrations in feather and tissue samples from hummingbirds and suggests a reference mark for exceeding normal. Lastly, pollinating avian species are minimally represented in the literature as bioindicators for environmental trace element contamination. Given that trace elements can move through food chains by a variety of routes, our study indicates that hummingbirds are possible bioindicators of environmental trace element contamination.

Winter range expansion of a hummingbird is associated with urbanization and supplementary feeding.

Anthropogenic changes to the landscape and climate cause novel ecological and evolutionary pressures, leading to potentially dramatic changes in the distribution of biodiversity. Warm winter temperatures can shift species' distributions to regions that were previously uninhabitable. Further, urbanization and supplementary feeding may facilitate range expansions and potentially reduce migration tendency. Here we explore how these factors interact to cause non-uniform effects across a species's range. Using 17 years of data from the citizen science programme Project FeederWatch, we examined the relationships between urbanization, winter temperatures and the availability of supplementary food (i.e. artificial nectar) on the winter range expansion (more than 700 km northward in the past two decades) of Anna's hummingbirds (Calypte anna). We found that Anna's hummingbirds have colonized colder locations over time, were more likely to colonize sites with higher housing density and were more likely to visit feeders in the expanded range compared to the historical range. Additionally, their range expansion mirrored a corresponding increase over time in the tendency of people to provide nectar feeders in the expanded range. This work illustrates how humans may alter the distribution and potentially the migratory behaviour of species through landscape and resource modification.

Neurons Responsive to Global Visual Motion Have Unique Tuning Properties in Hummingbirds

Neurons in animal visual systems that respond to global optic flow exhibit selectivity for motion direction and/or velocity. The avian lentiformis mesencephali (LM), known in mammals as the nucleus of the optic tract (NOT), is a key nucleus for global motion processing [1-4]. In all animals tested, it has been found that the majority of LM and NOT neurons aretuned to temporo-nasal (back-to-front) motion [4-11]. Moreover, the monocular gain of the optokinetic response is higher in this direction, compared to naso-temporal (front-to-back) motion [12, 13]. Hummingbirds are sensitive to small visual perturbations while hovering, and they drift to compensate for optic flow in all directions [14]. Interestingly, the LM, but not other visual nuclei, is hypertrophied in hummingbirds relative to other birds [15], which suggests enhanced perception of global visual motion. Using extracellular recording techniques, we found that there is a uniform distribution of preferred directions in the LM in Anna's hummingbirds, whereas zebra finch and pigeon LM populations, as in other tetrapods, show a strong bias toward temporo-nasal motion. Furthermore, LM and NOT neurons are generally classified as tuned to "fast" or "slow" motion [10, 16, 17], and we predicted that most neurons would be tuned to slow visual motion as an adaptation for slow hovering. However, we found the opposite result: most hummingbird LM neurons are tuned to fast pattern velocities, compared to zebra finches and pigeons. Collectively, these results suggest a role in rapid responses during hovering, as well as in velocity control and collision avoidance during forward flight of hummingbirds.

On the autorotation of animal wings.

Botanical samaras spin about their centre of mass and create vertical aerodynamic forces which slow their rate of descent. Descending autorotation of animal wings, however, has never been documented. We report here that isolated wings from Anna's hummingbirds, and also from 10 species of insects, can stably autorotate and achieve descent speeds and aerodynamic performance comparable to those of samaras. A hummingbird wing loaded at its base with the equivalent of 50% of the bird's body mass descended only twice as fast as an unloaded wing, and rotated at frequencies similar to those of the wings in flapping flight. We found that even entire dead insects could stably autorotate depending on their wing postures. Feather removal trials showed no effect on descent velocity when the secondary feathers were removed from hummingbird wings. By contrast, partial removal of wing primaries substantially improved performance, except when only the outer primary was present. A scaling law for the aerodynamic performance of autorotating wings is well supported if the wing aspect ratio and the relative position of the spinning axis from the wing base are included. Autorotation is a useful and practical method that can be used to explore the aerodynamics of wing design.

Mechanical Constraints on Flight at High Elevation Decrease Maneuvering Performance of Hummingbirds

High-elevation habitats offer ecological advantages including reduced competition, predation, and parasitism [1]. However, flying organisms at high elevation also face physiological challenges due to lower air density and oxygen availability [2]. These constraints are expected to affect the flight maneuvers that are required to compete with rivals, capture prey, and evade threats [3-5]. To test how individual maneuvering performance is affected by elevation, we measured the free-flight maneuvers of male Anna's hummingbirds in a large chamber translocated to a high-elevation site and then measured their performance at low elevation. We used a multi-camera tracking system to identify thousands of maneuvers based on body position and orientation [6]. At high elevation, the birds' translational velocities, accelerations, and rotational velocities were reduced, and they used less demanding turns. To determine how mechanical and metabolic constraints independently affect performance, we performed a second experiment to evaluate flight maneuvers in an airtight chamber infused with either normoxic heliox, to lower air density, or nitrogen, to lower oxygen availability. The hypodense treatment caused the birds to reduce their accelerations and rotational velocities, whereas the hypoxic treatment had no significant effect on maneuvering performance. Collectively, these experiments reveal how aerial maneuvering performance changes with elevation, demonstrating that as birds move up in elevation, air density constrains their maneuverability prior to any influence of oxygen availability. Our results support the hypothesis that changes in competitive ability at high elevations are the result of mechanical limits to flight performance [7].

Burst muscle performance predicts the speed, acceleration, and turning performance of Anna's hummingbirds.

Despite recent advances in the study of animal flight, the biomechanical determinants of maneuverability are poorly understood. It is thought that maneuverability may be influenced by intrinsic body mass and wing morphology, and by physiological muscle capacity, but this hypothesis has not yet been evaluated because it requires tracking a large number of free flight maneuvers from known individuals. We used an automated tracking system to record flight sequences from 20 Anna's hummingbirds flying solo and in competition in a large chamber. We found that burst muscle capacity predicted most performance metrics. Hummingbirds with higher burst capacity flew with faster velocities, accelerations, and rotations, and they used more demanding complex turns. In contrast, body mass did not predict variation in maneuvering performance, and wing morphology predicted only the use of arcing turns and high centripetal accelerations. Collectively, our results indicate that burst muscle capacity is a key predictor of maneuverability.

Electrostatic Charge on Flying Hummingbirds and Its Potential Role in Pollination.

Electrostatic phenomena are known to enhance both wind- and insect-mediated pollination, but have not yet been described for nectar-feeding vertebrates. Here we demonstrate that wild Anna's Hummingbirds (Calypte anna) can carry positive charges up to 800 pC while in flight (mean ± s.d.: 66 ± 129 pC). Triboelectric charging obtained by rubbing an isolated hummingbird wing against various plant structures generated charges up to 700 pC. A metal hummingbird model charged to 400 pC induced bending of floral stamens in four plants (Nicotiana, Hemerocallis, Penstemon, and Aloe spp.), and also attracted falling Lycopodium spores at distances of < 2 mm. Electrostatic forces may therefore influence pollen transfer onto nectar-feeding birds.

Hovering performance of Anna's hummingbirds (Calypte anna) in ground effect.

Aerodynamic performance and energetic savings for flight in ground effect are theoretically maximized during hovering, but have never been directly measured for flying animals. We evaluated flight kinematics, metabolic rates and induced flow velocities for Anna's hummingbirds hovering at heights (relative to wing length R = 5.5 cm) of 0.7R, 0.9R, 1.1R, 1.7R, 2.2R and 8R above a solid surface. Flight at heights less than or equal to 1.1R resulted in significant reductions in the body angle, tail angle, anatomical stroke plane angle, wake-induced velocity, and mechanical and metabolic power expenditures when compared with flight at the control height of 8R. By contrast, stroke plane angle relative to horizontal, wingbeat amplitude and wingbeat frequency were unexpectedly independent of height from ground. Qualitative smoke visualizations suggest that each wing generates a vortex ring during both down- and upstroke. These rings expand upon reaching the ground and present a complex turbulent interaction below the bird's body. Nonetheless, hovering near surfaces results in substantial energetic benefits for hummingbirds, and by inference for all volant taxa that either feed at flowers or otherwise fly close to plant or other surfaces.

Into turbulent air: size-dependent effects of von Kármán vortex streets on hummingbird flight kinematics and energetics

Animal fliers frequently move through a variety of perturbed flows during their daily aerial routines. However, the extent to which these perturbations influence flight control and energetic expenditure is essentially unknown. Here, we evaluate the kinematic and metabolic consequences of flight within variably sized vortex shedding flows using five Anna's hummingbirds feeding from an artificial flower in steady control flow and within vortex wakes produced behind vertical cylinders. Tests were conducted at three horizontal airspeeds (3, 6 and 9 m s(-1)) and using three different wake-generating cylinders (with diameters equal to 38, 77 and 173% of birds' wing length). Only minimal effects on wing and body kinematics were demonstrated for flight behind the smallest cylinder, whereas flight behind the medium-sized cylinder resulted in significant increases in the variances of wingbeat frequency, and variances of body orientation, especially at higher airspeeds. Metabolic rate was, however, unchanged relative to that of unperturbed flight. Hummingbirds flying within the vortex street behind the largest cylinder exhibited highest increases in variances of wingbeat frequency, and of body roll, pitch and yaw amplitudes at all measured airspeeds. Impressively, metabolic rate under this last condition increased by up to 25% compared with control flights. Cylinder wakes sufficiently large to interact with both wings can thus strongly affect stability in flight, eliciting compensatory kinematic changes with a consequent increase in flight metabolic costs. Our findings suggest that vortical flows frequently encountered by aerial taxa in diverse environments may impose substantial energetic costs.

Structure of the vortex wake in hovering Anna's hummingbirds (Calypte anna)

Hummingbirds are specialized hoverers for which the vortex wake has been described as a series of single vortex rings shed primarily during the downstroke. Recent findings in bats and birds, as well as in a recent study on Anna's hummingbirds, suggest that each wing may shed a discrete vortex ring, yielding a bilaterally paired wake. Here, we describe the presence of two discrete rings in the wake of hovering Anna's hummingbirds, and also infer force production through a wingbeat with contributions to weight support. Using flow visualization, we found separate vortices at the tip and root of each wing, with 15% stronger circulation at the wingtip than at the root during the downstroke. The upstroke wake is more complex, with near-continuous shedding of vorticity, and circulation of approximately equal magnitude at tip and root. Force estimates suggest that the downstroke contributes 66% of required weight support, whereas the upstroke generates 35%. We also identified a secondary vortex structure yielding 8-26% of weight support. Lift production in Anna's hummingbirds is more evenly distributed between the stroke phases than previously estimated for Rufous hummingbirds, in accordance with the generally symmetric down- and upstrokes that characterize hovering in these birds.

Implications of floral orientation for flight kinematics and metabolic expenditure of hover‐feeding hummingbirds

Summary Nectar‐bearing flowers are characterized by many different shapes, sizes and orientations, which may affect the way hummingbirds feed from them. Many hummingbird‐pollinated flowers are oriented downwards, thereby requiring that trochilids feed while hovering with the bill oriented vertically upward. We measured body orientations, wingbeat kinematics and hovering metabolic rates for Anna's Hummingbirds (Calypte anna) feeding from artificial flowers that were oriented horizontally, tilted 45° downwards and pointing vertically downwards. When feeding from vertically oriented flowers, hummingbirds employed an upright body position combined with dorsal head flexion. Additional kinematic adjustments included an increased stroke plane angle relative to the longitudinal body axis and an increased stroke amplitude deriving from increases in the minimum positional angle of the wingbeat. By contrast, wingbeat frequency, the stroke plane angle relative to horizontal, the ratio of the minimum to maximum positional angles of the wingbeat and the upstroke/downstroke ratio did not vary during feeding from different flower orientations. Metabolic rates increased by an average (±SD) of 10·8 (±8·8)% for feeding from vertically compared to horizontally oriented flowers. Feeding from pendent flowers comes with a substantial metabolic cost that may influence floral selection by hummingbirds and thus the evolution of associated pollination syndromes.

HUMMINGBIRDS MAKE FLYING BACKWARDS LOOK EASY

Backing up usually isn't easy, yet when Nir Sapir observed agile hummingbirds visiting a feeder on his balcony in Berkeley, California, he was struck by their ability to reverse. ‘I saw that they quite often fly backwards’, he recalls, adding that they always reverse out of a bloom after feasting. However, when he searched the literature he was disappointed to find that there were hardly any studies of this particular behaviour. ‘This was a bit surprising given that they are doing this all the time’, Sapir says, explaining that the tiny aviators visit flowers to feed once every 2–10 min. ‘I thought that this was an interesting topic to learn how they are doing it and what the consequences are for their metabolism’, Sapir says, so he and his postdoc advisor, Robert Dudley, set about measuring the flight movements and metabolism of reversing hummingbirds (p. 3603).Capturing five Anna's hummingbirds at a feeder located just inside a University of California Berkeley laboratory window, Sapir trained the birds to fly in a wind tunnel by tricking the birds into feeding from a syringe of sucrose disguised as a flower. He then filmed each bird as it hovered to feed before returning to the perch when satisfied. Knowing that the bird would return to the feeder again soon, Sapir turned on the air flow when the hummingbird arrived, directing the 3 m s−1 flow so that the bird had to fly backwards against the wind to remain stationary at the ‘flower’. Then he repeated the experiment with the syringe feeder rotated through 180 deg while the hummingbird flew forward into the wind to stay in place.Analysing the three flight styles, Sapir recalls that there were clear differences between forward and backward flight. The hummingbirds' body posture became much more upright as they flew backward, forcing them to bend their heads more to insert their beaks into the simulated flower. In addition, the reversing birds reduced the inclination of the plane of the wing beat so that it became more horizontal. And when Sapir analysed the wing beat frequency, he found that the birds were beating their wings at 43.8 Hz, instead of the 39.7 Hz that they use while flying forward. ‘That is quite a lot for hummingbirds because they hardly change their wing beat frequency’, explains Sapir.Repeating the experiments while recording the birds' oxygen consumption rates, Sapir says, ‘We expected that we would find high or intermediate values for metabolism during backward flight because the bird has an upright body position and this means that they have a higher drag. Also, the birds use backward flight frequently, but not all the time, so we assumed that it would not be more efficient in terms of the flight mechanics compared with forward flight.’ However, Sapir was surprised to discover that instead of being more costly, backward flight was as cheap as forward flight and 20% more efficient than hovering. And when Sapir gently increased the wind flow from 0 m s−1 in 1.5 m s−1 steps he found that their flight was cheapest at speeds of 3 m s−1 and above, although the birds were unable to fly backwards above 4.5 m s−1.Describing hummingbirds as insects trapped in a bird's body, Sapir adds that the fluttering flight of hummingbirds has more in common with insects than with their feathered cousins and he is keen to find out whether other hovering animals such as small songbirds and nectar-feeding bats can reverse too.

Backward flight in hummingbirds employs unique kinematic adjustments and entails low metabolic cost

Backward flight is a frequently used transient flight behavior among members of the species-rich hummingbird family (Trochilidae) when retreating from flowers, and is known from a variety of other avian and hexapod taxa, but the biomechanics of this intriguing locomotor mode have not been described. We measured rates of oxygen uptake (V(O2)) and flight kinematics of Anna's hummingbirds, Calypte anna (Lesson), within a wind tunnel using mask respirometry and high-speed videography, respectively, during backward, forward and hovering flight. We unexpectedly found that in sustained backward flight is similar to that in forward flight at equivalent airspeed, and is about 20% lower than hovering V(O2). For a bird that was measured throughout a range of backward airspeeds up to a speed of 4.5 m s(-1), the power curve resembled that of forward flight at equivalent airspeeds. Backward flight was facilitated by steep body angles coupled with substantial head flexion, and was also characterized by a higher wingbeat frequency, a flat stroke plane angle relative to horizontal, a high stroke plane angle relative to the longitudinal body axis, a high ratio of maximum:minimum wing positional angle, and a high upstroke:downstroke duration ratio. Because of the convergent evolution of hummingbird and some hexapod flight styles, flying insects may employ similar kinematics while engaged in backward flight, for example during station keeping or load lifting. We propose that backward flight behavior in retreat from flowers, together with other anatomical, physiological, morphological and behavioral adaptations, enables hummingbirds to maintain strictly aerial nectarivory.

Dietary protein level affects iridescent coloration in Anna's hummingbirds, Calypte anna

Many animal displays involve colorful ornamental traits that signal an individual's quality as a mate or rival. Brilliant iridescent ornaments are common, but little is currently known about their production cost and signaling value. One potential cost of colorful ornaments is the acquisition of limited dietary resources that may be involved, directly or indirectly, in their production. Protein, the primary component of bird feathers and of many nanostructural components of iridescent traits, is naturally restricted in hummingbird diets (comprised mostly of sugars), suggesting that iridescent coloration may be especially challenging to produce in these animals. In this study, we experimentally investigated the effect of dietary protein availability during molt on iridescent color expression in male Anna's hummingbirds (Calypte anna). We fed captive birds either a 6% (high) or a 3% (low) protein diet and stimulated molt by plucking half the gorget and crown ornaments on each bird as well as the non-ornamental iridescent green tail feathers. We found that birds receiving more protein grew significantly more colorful crown feathers (higher red chroma and redder hue) than those fed the low-protein diet. Diet did not affect gorget coloration, but regrowth of feathers in captivity affected both gorget and crown coloration. Additionally, birds on the high-protein diet grew yellower (higher hue) green tail feathers than birds on the low-protein diet. These results indicate that iridescent ornamental feathers are sensitive to diet quality and may serve as honest signals of nutrition to mates or rivals. Further, because both ornamental and non-ornamental iridescent coloration were affected by conditions during their growth, iridescent color in these birds appears to be generally condition dependent.

Flying in the rain: hovering performance of Anna's hummingbirds under varied precipitation

Flight in rain represents a greater challenge for smaller animals because the relative effects of water loading and drop impact are greater at reduced scales given the increased ratios of surface area to mass. Nevertheless, it is well known that small volant taxa such as hummingbirds can continue foraging even in extreme precipitation. Here, we evaluated the effect of four rain intensities (i.e. zero, light, moderate and heavy) on the hovering performance of Anna's hummingbirds (Calypte anna) under laboratory conditions. Light-to-moderate rain had only a marginal effect on flight kinematics; wingbeat frequency of individuals in moderate rain was reduced by 7 per cent relative to control conditions. By contrast, birds hovering in heavy rain adopted more horizontal body and tail positions, and also increased wingbeat frequency substantially, while reducing stroke amplitude when compared with control conditions. The ratio between peak forces produced by single drops on a wing and on a solid surface suggests that feathers can absorb associated impact forces by up to approximately 50 per cent. Remarkably, hummingbirds hovered well even under heavy precipitation (i.e. 270 mm h(-1)) with no apparent loss of control, although mechanical power output assuming perfect and zero storage of elastic energy was estimated to be about 9 and 57 per cent higher, respectively, compared with normal hovering.

Corrigendum for the paper ‘Aerial shaking performance of wet Anna's hummingbirds’

[ J. R. Soc. Interface 9 , 1093–1099 (2012; Published online 9 November 2011) ([doi:10.1098/rsif.2011.0608][2])][2] In our recently published article, we stated in the Abstract that ‘Tangential speeds and accelerations of the trunk and tail were roughly similar in aerial and non-aerial shakes,

Wingbeat kinematics and motor control of yaw turns in Anna's hummingbirds (Calypte anna)

The biomechanical and neuromuscular mechanisms used by different animals to generate turns in flight are highly variable. Body size and body plan exert some influence, e.g. birds typically roll their body to orient forces generated by the wings whereas insects are capable of turning via left-right wingbeat asymmetries. Turns are also relatively brief and have low repeatability, with almost every wingbeat serving a different function throughout the change in heading. Here we present an analysis of Anna's hummingbirds (Calypte anna) as they fed continuously from an artificial feeder revolving around the outside of the animal. This setup allowed for examination of sustained changes in yaw without requiring any corresponding changes in pitch, roll or body position. Hummingbirds sustained yaw turns by expanding the wing stroke amplitude of the outer wing during the downstroke and by altering the deviation of the wingtip path during both downstroke and upstroke. The latter led to a shift in the inner-outer stroke plane angle during the upstroke and shifts in the elevation of the stroke plane and in the deviation of the wingtip path during both strokes. These features are generally more similar to how insects, as opposed to birds, turn. However, time series analysis also revealed considerable stroke-to-stroke variation. Changes in the stroke amplitude and the wingtip velocity were highly cross-correlated, as were changes in the stroke deviation and the elevation of the stroke plane. As was the case for wingbeat kinematics, electromyogram recordings from pectoral and wing muscles were highly variable, but no correlations were found between these two features of motor control. The high variability of both kinematic and muscle activation features indicates a high level of wingbeat-to-wingbeat adjustments during sustained yaw. The activation timing of the muscles was more repeatable than the activation intensity, which suggests that the former may be constrained by harmonic motion and that the latter may play a large role in kinematic adjustments. Comparing the revolution frequency of the feeder with measurements of free flight yaws reveals that feeder tracking, even at one revolution every 2 s, is well below the maximum yaw capacity of the hummingbirds.

Aerial shaking performance of wet Anna's hummingbirds

External wetting poses problems of immediate heat loss and long-term pathogen growth for vertebrates. Beyond these risks, the locomotor ability of smaller animals, and particularly of fliers, may be impaired by water adhering to the body. Here, we report on the remarkable ability of hummingbirds to perform rapid shakes in order to expel water from their plumage even while in flight. Kinematic performance of aerial versus non-aerial shakes (i.e. those performed while perching) was compared. Oscillation frequencies of the head, body and tail were lower in aerial shakes. Tangential speeds and accelerations of the trunk and tail were roughly similar in aerial and non-aerial shakes, but values for head motions while perching were twice as high when compared with aerial shakes [corrected] . Azimuthal angular amplitudes for both aerial and non-aerial shakes reached values greater than 180° for the head, greater than 45° for the body trunk and slightly greater than 90° for the tail and wings. Using a feather on an oscillating disc to mimic shaking motions, we found that bending increased average speeds by up to 36 per cent and accelerations of the feather tip up to fourfold relative to a hypothetical rigid feather. Feather flexibility may help to enhance shedding of water and reduce body oscillations during shaking.

Hovering and Forward Flight Energetics in Anna's and Allen's Hummingbirds

Aerodynamic theory predicts that the mechanical costs of flight are lowest at intermediate flight speeds; metabolic costs of flight should trend similarly if muscle efficiency is constant. We measured metabolic rates for nine Anna's hummingbirds (Calypte anna) and two male Allen's hummingbirds (Selasphorus sasin) feeding during flight from a free-standing mask over a range of airspeeds. Ten of 11 birds exhibited higher metabolic costs during hovering than during flight at intermediate airspeeds, whereas one individual exhibited comparable costs at hovering and during forward flight up to speeds of approximately 7 m s(-1). Flight costs of all hummingbirds increased at higher airspeeds. Relative to Anna's hummingbirds, Allen's hummingbirds exhibited deeper minima in the power curve, possibly due to higher wing loadings and greater associated costs of induced drag. Although feeding at a mask in an airstream may reduce body drag and, thus, the contributions of parasite power to overall metabolic expenditure, these results suggest that hummingbird power curves are characterized by energetic minima at intermediate speeds relative to hovering costs.