Flight Style Research Articles

The influence of flight style on the aerodynamic properties of avian wings as fixed lifting surfaces.

The diversity of wing morphologies in birds reflects their variety of flight styles and the associated aerodynamic and inertial requirements. Although the aerodynamics underlying wing morphology can be informed by aeronautical research, important differences exist between planes and birds. In particular, birds operate at lower, transitional Reynolds numbers than do most aircraft. To date, few quantitative studies have investigated the aerodynamic performance of avian wings as fixed lifting surfaces and none have focused upon the differences between wings from different flight style groups. Dried wings from 10 bird species representing three distinct flight style groups were mounted on a force/torque sensor within a wind tunnel in order to test the hypothesis that wing morphologies associated with different flight styles exhibit different aerodynamic properties. Morphological differences manifested primarily as differences in drag rather than lift. Maximum lift coefficients did not differ between groups, whereas minimum drag coefficients were lowest in undulating flyers (Corvids). The lift to drag ratios were lower than in conventional aerofoils and data from free-flying soaring species; particularly in high frequency, flapping flyers (Anseriformes), which do not rely heavily on glide performance. The results illustrate important aerodynamic differences between the wings of different flight style groups that cannot be explained solely by simple wing-shape measures. Taken at face value, the results also suggest that wing-shape is linked principally to changes in aerodynamic drag, but, of course, it is aerodynamics during flapping and not gliding that is likely to be the primary driver.

Evolution of avian flight: muscles and constraints on performance.

Competing hypotheses about evolutionary origins of flight are the 'fundamental wing-stroke' and 'directed aerial descent' hypotheses. Support for the fundamental wing-stroke hypothesis is that extant birds use flapping of their wings to climb even before they are able to fly; there are no reported examples of incrementally increasing use of wing movements in gliding transitioning to flapping. An open question is whether locomotor styles must evolve initially for efficiency or if they might instead arrive due to efficacy. The proximal muscles of the avian wing output work and power for flight, and new research is exploring functions of the distal muscles in relation to dynamic changes in wing shape. It will be useful to test the relative contributions of the muscles of the forearm compared with inertial and aerodynamic loading of the wing upon dynamic morphing. Body size has dramatic effects upon flight performance. New research has revealed that mass-specific muscle power declines with increasing body mass among species. This explains the constraints associated with being large. Hummingbirds are the only species that can sustain hovering. Their ability to generate force, work and power appears to be limited by time for activation and deactivation within their wingbeats of high frequency. Most small birds use flap-bounding flight, and this flight style may offer an energetic advantage over continuous flapping during fast flight or during flight into a headwind. The use of flap-bounding during slow flight remains enigmatic. Flap-bounding birds do not appear to be constrained to use their primary flight muscles in a fixed manner. To improve understanding of the functional significance of flap-bounding, the energetic costs and the relative use of alternative styles by a given species in nature merit study.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'.

Physiological, aerodynamic and geometric constraints of flapping account for bird gaits, and bounding and flap-gliding flight strategies

Aerodynamically economical flight is steady and level. The high-amplitude flapping and bounding flight style of many small birds departs considerably from any aerodynamic or purely mechanical optimum. Further, many large birds adopt a flap-glide flight style in cruising flight which is not consistent with purely aerodynamic economy. Here, an account is made for such strategies by noting a well-described, general, physiological cost parameter of muscle: the cost of activation. Small birds, with brief downstrokes, experience disproportionately high costs due to muscle activation for power during contraction as opposed to work. Bounding flight may be an adaptation to modulate mean aerodynamic force production in response to (1) physiological pressure to extend the duration of downstroke to reduce power demands during contraction; (2) the prevention of a low-speed downstroke due to the geometric constraints of producing thrust; (3) an aerodynamic cost to flapping with very low lift coefficients. In contrast, flap-gliding birds, which tend to be larger, adopt a strategy that reduces the physiological cost of work due both to activation and contraction efficiency. Flap-gliding allows, despite constraints to modulation of aerodynamic force lever-arm, (1) adoption of moderately large wing-stroke amplitudes to achieve suitable muscle strains, thereby reducing the activation costs for work; (2) reasonably quick downstrokes, enabling muscle contraction at efficient velocities, while being (3) prevented from very slow weight-supporting upstrokes due to the cost of performing ‘negative’ muscle work.

Multitrait aposematic signal in Batesian mimicry

Batesian mimics can parasitize Müllerian mimicry rings mimicking the warning color signal. The evolutionary success of Batesian mimics can increase adding complexity to the signal by behavioral and locomotor mimicry. We investigated three fundamental morphological and locomotor traits in a Neotropical mimicry ring based on Ithomiini butterflies and parasitized by Polythoridae damselflies: wing color, wing shape, and flight style. The study species have wings with a subapical white patch, considered the aposematic signal, and a more apical black patch. The main predators are VS-birds, visually more sensitive to violet than to ultraviolet wavelengths (UVS-birds). The white patches, compared to the black patches, were closer in the bird color space, with higher overlap for VS-birds than for UVS-birds. Using a discriminability index for bird vision, the white patches were more similar between the mimics and the model than the black patches. The wing shape of the mimics was closer to the model in the morphospace, compared to other outgroup damselflies. The wing-beat frequency was similar among mimics and the model, and different from another outgroup damselfly. Multitrait aposematic signals involving morphology and locomotion may favor the evolution of mimicry rings and the success of Batesian mimics by improving signal effectiveness toward predators.

Predictable evolution toward flightlessness in volant island birds

Birds are prolific colonists of islands, where they readily evolve distinct forms. Identifying predictable, directional patterns of evolutionary change in island birds, however, has proved challenging. The "island rule" predicts that island species evolve toward intermediate sizes, but its general applicability to birds is questionable. However, convergent evolution has clearly occurred in the island bird lineages that have undergone transitions to secondary flightlessness, a process involving drastic reduction of the flight muscles and enlargement of the hindlimbs. Here, we investigated whether volant island bird populations tend to change shape in a way that converges subtly on the flightless form. We found that island bird species have evolved smaller flight muscles than their continental relatives. Furthermore, in 366 populations of Caribbean and Pacific birds, smaller flight muscles and longer legs evolved in response to increasing insularity and, strikingly, the scarcity of avian and mammalian predators. On smaller islands with fewer predators, birds exhibited shifts in investment from forelimbs to hindlimbs that were qualitatively similar to anatomical rearrangements observed in flightless birds. These findings suggest that island bird populations tend to evolve on a trajectory toward flightlessness, even if most remain volant. This pattern was consistent across nine families and four orders that vary in lifestyle, foraging behavior, flight style, and body size. These predictable shifts in avian morphology may reduce the physical capacity for escape via flight and diminish the potential for small-island taxa to diversify via dispersal.

Anatomical differences in three wing muscles of the Grey heron (Ardea cinerea), the Common buzzard (Buteo buteo) and the Common kestrel (Falco tinnunculus): a possible functional interpretation

In this paper, a description of the coracobrachialis caudalis, the scapulotriceps and the extensor carpi radialis muscles is presented in three different species (Common kestrel, Common buzzard and Grey heron) that exhibit notably differing flight styles. The primary goal of this research is to describe the gross anatomy of species not previously examined. Secondly, we attempt to advance a functional interpretation of the structures studied, to understand if such noticeable differences in flight style could have induced a differentiation in the above mentioned wing muscles. Regarding the coracobrachialis caudalis muscle, the Grey heron exhibited an unusual conformation with two “reversed” heads and a great amount of fibrous bundles. Moreover, the “anchors” of the scapulotriceps muscle were well developed in the Grey heron and vestigial in the Common kestrel, while an intermediate situation was observed in the Common buzzard. In addition, the extensor carpi radialis muscle showed interesting findings concerning the number of heads and their disposition. Our data may corroborate some previously advanced functional interpretation. In particular, the structure of the coracobrachialis muscle is in line with the hypothesis of its role as muscular strut. Moreover, the humeral and scapular anchors could play a role in increasing the scapulotriceps tension during some phases of the beat cycle or it may be involved in maintaining wing posture. Regarding the different number of heads and the disposition of the extensor carpi radialis muscle, we believe that it could play a role in maintaining the wing position. Although a correct functional interpretation can be obtained only through multidisciplinary studies, we believe that an in-depth gross anatomy study should always be conducted beforehand, to serve as basis for further and more specific research.

The evolution of avian wing shape and previously unrecognized trends in covert feathering.

Avian wing shape has been related to flight performance, migration, foraging behaviour and display. Historically, linear measurements of the feathered aerofoil and skeletal proportions have been used to describe this shape. While the distribution of covert feathers, layered over the anterior wing, has long been assumed to contribute to aerofoil properties, to our knowledge no previous studies of trends in avian wing shape assessed their variation. Here, these trends are explored using a geometric-morphometric approach with landmarks describing the wing outline as well as the extent of dorsal and ventral covert feathers for 105 avian species. We find that most of the observed variation is explained by phylogeny and ecology but shows only a weak relationship with previously described flight style categories, wing loading and an investigated set of aerodynamic variables. Most of the recovered variation is in greater primary covert feather extent, followed by secondary feather length and the shape of the wing tip. Although often considered a plastic character strongly linked to flight style, the estimated ancestral wing morphology is found to be generally conservative among basal parts of most major avian lineages. The radiation of birds is characterized by successive diversification into largely distinct areas of morphospace. However, aquatic taxa show convergence in feathering despite differences in flight style, and songbirds move into a region of morphospace also occupied by basal taxa but at markedly different body sizes. These results have implications for the proposed inference of flight style in extinct taxa.

Signalling with clear wings during territorial behaviour and courtship ofChlorocypha cancellata(Odonata, Chlorocyphidae)

The reproductive behaviour of the damselfly Chlorocypha cancellata (Chlorocyphidae) was filmed at 600 frames per second. Different flight styles including straight, forward flight, threat and courtship display were analysed with respect to changes in wing beat frequency and phase relationships of fore and hind wings. The analysis revealed significant differences in the flight style between non-escalated and escalated threat display as well as changes in the wing beat frequency of a male during courtship dependant on the behaviour of the female. This is the first evidence suggesting that odonate species with clear wings can use specialized flight modes for intraspecific signalling.

Predictive Effect of Implicit Safety Attitudes on Safety Performance in Aviation Safety Culture

The fundamental feature of safety culture is represented by safety attitudes. In terms of measuring safety cultures, many researchers have mainly focused on explicit safety attitudes and generally relied on specific survey instruments. It is questionable, however, whether self-report measures can capture all aspects of organizational safety culture. Instead of getting direct answers, it is necessary to introduce implicit measures and the implicit safety attitudes test into safety culture evaluation. The basic hypothesis of this study was that while various enterprises share different safety culture, the structure and intensity of implicit and explicit safety attitudes differ for employees. The present study was aimed at investigating the complete model of aviation safety culture and the importance of implicit safety attitudes by detecting the relationship between explicit and implicit safety attitudes as well as the prediction effect of implicit safety attitudes.The Flight Management Attitudes Questionnaire(FMAQ 2.0, international version) was adopted in this study which was based on the work characteristics of modern airlines pilots. With the purpose of evaluating explicit safety attitudes under the background of aviation safety culture, FMAQ 2.0 is comprised of three subscales, including basic organizational attitudes, cockpit work attitudes, and flight automation attitudes. Moreover, Evaluative Implicit Association Test and Affective Implicit Association Test were developed for aviatic implicit safety attitudes test. 134 pilots were involved in the investigation, 126 valid cases were obtained. Safety performance were obtained from airline company on four dimensions, including safety regulation, flight style, flight skill and organizational management was applied for validity criterion. The results showed that(1) IAT of aviation safety attitudes indicated a high effect value, aviation safety led to more positive evaluation and feelings while flight risk and adventure were more connected with negative evaluation and emotion.(2) Implicit and explicit safety attitudes were both relevant and relative separation. There was a positive relation between evaluative implicit safety attitudes and cockpit work attitudes. Affective implicit safety attitudes showed significant positive correlation with cockpit work attitudes and flight automation attitudes. And the model data showed that implicit and explicit safety attitudes were separation structure.(3) Both the flight management attitudes and implicit safety attitudes were able to predict safety performance. The former were doing better on predicting flight style, flight skill and organizational management, while the latter had higher prediction rate on safety regulation. The common prediction model fitted well. The study demonstrated a high intensity of aviation safety attitudes. IAT can be used as an effective evaluation tool for implicit safety attitudes. Explicit and implicit safety attitudes are both unified and separated structure. The explicit safety attitudes, implicit safety attitudes and safety performance together constitute a complete model of aviation safety culture. And in this model, the outer layer is aviation safety performance, the middle layer is explicit safety attitudes which can be measured directly, the inner layer is implicit safety attitudes which can be evaluated by indirect measurement methods.

Nanomechanical properties of bird feather rachises: exploring naturally occurring fibre reinforced laminar composites.

Flight feathers have evolved under selective pressures to be sufficiently light and strong enough to cope with the stresses of flight. The feather shaft (rachis) must resist these stresses and is fundamental to this mode of locomotion. Relatively little work has been done on rachis morphology, especially from a mechanical perspective and never at the nanoscale. Nano-indentation is a cornerstone technique in materials testing. Here we use this technique to make use of differentially oriented fibres and their resulting mechanical anisotropy. The rachis is established as a multi-layered fibrous composite material with varying laminar properties in three feathers of birds with markedly different flight styles; the Mute Swan (Cygnus olor), the Bald Eagle (Haliaeetus leucocephalus) and the partridge (Perdix perdix). These birds were chosen not just because they are from different clades and have different flight styles, but because they have feathers large enough to gain meaningful results from nano-indentation. Results from our initial datasets indicate that the proportions and orientation of the laminae are not fixed and may vary either in order to cope with the stresses of flight particular to the bird or with phylogenetic lineage.

Evolution of extreme proboscis lengths in Neotropical Hesperiidae (Lepidoptera)

Exaggerated morphologies have evolved in insects as adaptations to nectar feeding by natural selection. For example, the suctorial mouthparts of butterflies enable these insects to gain access to floral nectar concealed inside deep floral tubes. Proboscis length in Lepidoptera is known to scale with body size, but whether extreme absolute proboscis lengths of nectar feeding butterflies result from a proportional or disproportional increase with body size that differs between phylogenetic lineages remains unknown. We surveyed the range of variation that occurs in scaling relationships between proboscis length and body size against a phylogenetic background among Costa Rican Hesperiidae. We obtained a new record holder for the longest proboscis in butterflies and showed that extremely long proboscides evolved at least three times independently within Neotropical Hesperiidae. We conclude that the evolution of extremely long proboscides results from allometric scaling with body size, as demonstrated in hawk moths. We hypothesize that constraints on the evolution of increasingly long butterfly proboscides may come from (1) the underlying scaling relationships, i.e., relative proboscis length, combined with the butterfly's flight style and flower-visiting behaviour and/or (2) developmental constraints during the pupal phase. Lastly, we discuss why butterflies did not evolve similar scaling relationships as hawk moths.

Microstructure and cross-sectional shape of limb bones in Great Horned Owls and Red-tailed Hawks: how do these features relate to differences in flight and hunting behavior?

The Red-tailed Hawk and Great Horned Owl are two species of raptor that are similar in body size, diet, and habitat. Both species use their hindlimbs during hunting, but differ in foot morphology, how they approach and immobilize prey, and the average size of prey captured. They also differ in primary flight style: the Red-tailed Hawk uses static soaring and the Great Horned Owl uses flap-gliding. The objectives of this study were to characterize the microstructure and cross-sectional shape of limb bones of these species and examine the relationship with flight and hunting behaviors. The mid-shaft of six limb bones from six individuals of each species was sampled. The degree of bone laminarity (proportion of circular primary vascular canals) and cross-sectional geometric parameters were calculated. In both species, the humerus and femur exhibited features that suggest high resistance to torsional loading, whereas the tibiotarsus and phalanges had a shape more likely to resist compression and bending in a specific plane. The femur of the Red-tailed Hawk exhibited higher laminarity and larger polar moment of area than that of the Great Horned Owl. The tibiotarsus was more elliptical than that of the Great Horned Owl. The hawk approaches prey from a more horizontal axis, takes prey of greater mass, and is more likely to pursue prey on the ground, which could potentially be causing more torsional loads on the femur and bending loads on the tibiotarsus. In addition, differences in polar moment of area of the phalanges between the species could relate to differences in foot morphology or digit length. The humerus and ulna of the flap-gliding Great Horned Owl are more elliptical than the static soaring Red-tailed Hawk, a shape that may better resist the bending loads associated with a larger amount of flapping.

Phylogeny and forelimb disparity in waterbirds.

Previous work has shown that the relative proportions of wing components (i.e., humerus, ulna, carpometacarpus) in birds are related to function and ecology, but these have rarely been investigated in a phylogenetic context. Waterbirds including "Pelecaniformes," Ciconiiformes, Procellariiformes, Sphenisciformes, and Gaviiformes form a highly supported clade and developed a great diversity of wing forms and foraging ecologies. In this study, forelimb disparity in the waterbird clade was assessed in a phylogenetic context. Phylogenetic signal was assessed via Pagel's lambda, Blomberg's K, and permutation tests. We find that different waterbird clades are clearly separated based on forelimb component proportions, which are significantly correlated with phylogeny but not with flight style. Most of the traditional contents of "Pelecaniformes" (e.g., pelicans, cormorants, and boobies) cluster with Ciconiiformes (herons and storks) and occupy a reduced morphospace. These taxa are closely related phylogenetically but exhibit a wide range of ecologies and flight styles. Procellariiformes (e.g., petrels, albatross, and shearwaters) occupy a wide range of morphospace, characterized primarily by variation in the relative length of carpometacarpus and ulna. Gaviiformes (loons) surprisingly occupy a wing morphospace closest to diving petrels and penguins. Whether this result may reflect wing proportions plesiomorphic for the waterbird clade or a functional signal is unclear. A Bayesian approach detecting significant rate shifts across phylogeny recovered two such shifts. At the base of the two sister clades Sphenisciformes + Procellariiformes, a shift to an increase evolutionary rate of change is inferred for the ulna and carpometacarpus. Thus, changes in wing shape begin prior to the loss of flight in the wing-propelled diving clade. Several shifts to slower rate of change are recovered within stem penguins.

Morphological properties of the last primaries, the tail feathers, and the alulae of Accipiter nisus, Columba livia, Falco peregrinus, and Falco tinnunculus.

We investigated the mechanical properties (Young's modulus, bending stiffness, barb separation forces) of the tenth primary of the wings, of the alulae and of the middle tail feathers of Falco peregrinus. For comparison, we also investigated the corresponding feathers in pigeons (Columba livia), kestrels (Falco tinnunculus), and sparrowhawks (Accipiter nisus). In all four species, the Young's moduli of the feathers ranged from 5.9 to 8.4 GPa. The feather shafts of F. peregrinus had the largest cross-sections and the highest specific bending stiffness. When normalized with respect to body mass, the specific bending stiffness of primary number 10 was highest in F. tinnunculus, while that of the alula was highest in A. nisus. In comparison, the specific bending stiffness, measured at the base of the tail feathers and in dorso-ventral bending direction, was much higher in F. peregrinus than in the other three species. This seems to correlate with the flight styles of the birds: F. tinnunculus hovers and its primaries might therefore withstand large mechanical forces. A. nisus has often to change its flight directions during hunting and perhaps needs its alulae for this maneuvers, and in F. peregrinus, the base of the tail feathers might need a high stiffness during breaking after diving.

Ecology and caudal skeletal morphology in birds: the convergent evolution of pygostyle shape in underwater foraging taxa.

Birds exhibit a specialized tail that serves as an integral part of the flight apparatus, supplementing the role of the wings in facilitating high performance aerial locomotion. The evolution of this function for the tail contributed to the diversification of birds by allowing them to utilize a wider range of flight behaviors and thus exploit a greater range of ecological niches. The shape of the wings and the tail feathers influence the aerodynamic properties of a bird. Accordingly, taxa that habitually utilize different flight behaviors are characterized by different flight apparatus morphologies. This study explores whether differences in flight behavior are also associated with variation in caudal vertebra and pygostyle morphology. Details of the tail skeleton were characterized in 51 Aequornithes and Charadriiformes species. Free caudal vertebral morphology was measured using linear metrics. Variation in pygostyle morphology was characterized using Elliptical Fourier Analysis, a geometric morphometric method for the analysis of outline shapes. Each taxon was categorized based on flight style (flap, flap-glide, dynamic soar, etc.) and foraging style (aerial, terrestrial, plunge dive, etc.). Phylogenetic MANOVAs and Flexible Discriminant Analyses were used to test whether caudal skeletal morphology can be used to predict flight behavior. Foraging style groups differ significantly in pygostyle shape, and pygostyle shape predicts foraging style with less than 4% misclassification error. Four distinct lineages of underwater foraging birds exhibit an elongate, straight pygostyle, whereas aerial and terrestrial birds are characterized by a short, dorsally deflected pygostyle. Convergent evolution of a common pygostyle phenotype in diving birds suggests that this morphology is related to the mechanical demands of using the tail as a rudder during underwater foraging. Thus, distinct locomotor behaviors influence not only feather attributes but also the underlying caudal skeleton, reinforcing the importance of the entire caudal locomotor module in avian ecological diversification.

Flight style optimization in ski jumping on normal, large, and ski flying hills

In V-style ski jumping, aerodynamic forces are predominant performance factors and athletes have to solve difficult optimization problems in parts of a second in order to obtain their jump length maximum and to keep the flight stable. Here, a comprehensive set of wind tunnel data was used for optimization studies based on Pontryagin′s minimum principle with both the angle of attack α and the body-ski angle β as controls. Various combinations of the constraints αmax and βmin(t) were analyzed in order to compare different optimization strategies. For the computer simulation studies, the Olympic hill profiles in Esto-Sadok, Russia (HS 106m, HS 140m), and in Harrachov, Czech Republic, host of the Ski Flying World Championships 2014 (HS 205m) were used. It is of high importance for ski jumping practice that various aerodynamic strategies, i.e. combinations of α- and β-time courses, can lead to similar jump lengths which enables athletes to win competitions using individual aerodynamic strategies. Optimization results also show that aerodynamic behavior has to be different at different hill sizes (HS). Optimized time courses of α and β using reduced drag and lift areas in order to mimic recent equipment regulations differed only in a negligible way. This indicates that optimization results presented here are not very sensitive to minor changes of the aerodynamic equipment features when similar jump length are obtained by using adequately higher in-run velocities. However, wind tunnel measurements with athletes including take-off and transition to stabilized flight, flight, and landing behavior would enable a more detailed understanding of individual flight style optimization.

Correction: Avian Wing Proportions and Flight Styles: First Step towards Predicting the Flight Modes of Mesozoic Birds

Correction: Avian Wing Proportions and Flight Styles: First Step towards Predicting the Flight Modes of Mesozoic Birds

Fossil evidence of wing shape in a stem relative of swifts and hummingbirds (Aves, Pan-Apodiformes).

A feathered specimen of a new species of Eocypselus from the Early Eocene Green River Formation of Wyoming provides insight into the wing morphology and ecology in an early part of the lineage leading to extant swifts and hummingbirds. Combined phylogenetic analysis of morphological and molecular data supports placement of Eocypselus outside the crown radiation of Apodiformes. The new specimen is the first described fossil of Pan-Apodiformes from the pre-Pleistocene of North America and the only reported stem taxon with informative feather preservation. Wing morphology of Eocypselus rowei sp. nov. is intermediate between the short wings of hummingbirds and the hyper-elongated wings of extant swifts, and shows neither modifications for the continuous gliding used by swifts nor modifications for the hovering flight style used by hummingbirds. Elongate hindlimb elements, particularly the pedal phalanges, also support stronger perching capabilities than are present in Apodiformes. The new species is the smallest bird yet described from the Green River Formation, and supports the hypothesis that a decrease in body size preceded flight specializations in Pan-Apodiformes. The specimen also provides the first instance of melanosome morphology preserved in association with skeletal remains from the Green River Formation.

Locomotory abilities and habitat of the Cretaceous bird Gansus yumenensis inferred from limb length proportions

The relative length proportions of the three bony elements of the pelvic (femur, tibiotarsus and tarsometatarsus) and pectoral (humerus, ulna and manus) limbs of the early Cretaceous bird Gansus yumenensis, a well-represented basal ornithuromorph from China, are investigated and compared to those of extant taxa. Ternary plots show that the pectoral limb length proportions of Gansus are most similar to Apodiformes (swifts and hummingbirds), which plot away from all other extant birds. In contrast, the pelvic limb length proportions of Gansus fall within the extant bird cluster and show similarities with the neornithine families Podicipedidae (grebes), Diomedeidae (albatross) and Phalacrocoracidae (cormorants). Although it does have some of the pelvic limb features of grebes and cormorants, the femur of Gansus is more gracile and is thus more consistent with an albatross-like shallow-diving mode of life than a strong foot-propelled diving movement pattern. The position of Gansus in pectoral limb ternary morphospace is largely due to its elongated manus. In contrast to apodiformes, where the humerus and ulna are short and robust, an adaptation, which provides a stiff wing for their demanding fast agile and hovering flight (respectively), the wing-bones of Gansus are slender, indicating a less vigorous flapping flight style. The suite of characters exhibited by Gansus mean it is difficult to completely interpret its likely ecology. Nevertheless, our analyses suggest that it is probable that this bird was both volant and capable of diving to some degree using either foot-propelled or, perhaps, both its wings and its feet for underwater locomotion.

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.