Flight Style Research Articles

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.

Primary feather lengths may not be important for inferring the flight styles of Mesozoic birds

Although many Mesozoic fossil birds have been found with primary feathers preserved, these structures have rarely been included in morphometric analyses. This is surprising because the flight feathers of modern birds can contribute approximately 50% of the total wing length, and so it would be assumed that their inclusion or exclusion would modify functional interpretations. Here we show, contrary to earlier work, that this may not be the case. Using forelimb measurements and primary feather lengths from Mesozoic birds, we constructed morphospaces for different clades, which we then compared with morphospaces constructed for extant taxa classified according to flight mode. Consistent with older work, our results indicate that among extant birds some functional flight groups can be distinguished on the basis of their body sizes and that variation in the relative proportions of the wing elements is conservative. Mesozoic birds, on the other hand, show variable proportions of wing bones, with primary feather length contribution to the wing reduced in the earlier diverging groups. We show that the diverse Mesozoic avian clade Enantiornithes overlaps substantially with extant taxa in both size and limb element proportions, confirming previous morphometric results based on skeletal elements alone. However, these measurements cannot be used to distinguish flight modes in extant birds, and so cannot be used to infer flight mode in fossil forms. Our analyses suggest that more data from fossil birds, combined with accurate functional determination of the flight styles of living forms is required if we are to be able to predict the flight modes of extinct birds

Functional morphometric analysis of the furcula in mesozoic birds.

The furcula displays enormous morphological and structural diversity. Acting as an important origin for flight muscles involved in the downstroke, the form of this element has been shown to vary with flight mode. This study seeks to clarify the strength of this form-function relationship through the use of eigenshape morphometric analysis coupled with recently developed phylogenetic comparative methods (PCMs), including phylogenetic Flexible Discriminant Analysis (pFDA). Additionally, the morphospace derived from the furculae of extant birds is used to shed light on possible flight adaptations of Mesozoic fossil taxa. While broad conclusions of earlier work are supported (U-shaped furculae are associated with soaring, strong anteroposterior curvature with wing-propelled diving), correlations between form and function do not appear to be so clear-cut, likely due to the significantly larger dataset and wider spectrum of flight modes sampled here. Interclavicular angle is an even more powerful discriminator of flight mode than curvature, and is positively correlated with body size. With the exception of the close relatives of modern birds, the ornithuromorphs, Mesozoic taxa tend to occupy unique regions of morphospace, and thus may have either evolved unfamiliar flight styles or have arrived at similar styles through divergent musculoskeletal configurations.

Eigensurface analysis, ecology, and modelling of morphological adaptation in the falconiform humerus (Falconiformes: Aves)

Studies of morphological adaptation aim to quantify the relationship between an organism's form and its ecology. In the past such studies have been hampered by an over-reliance on either qualitative observations or the collection of a few, marginally representative two-dimensional linear measurements as morphological descriptors. Recent advances in morphometric data acquisition and analysis techniques now provide a means of accurately and comprehensively quantifying the morphological variation inherent in complex three-dimensional (3D) surfaces across a specimen set. Using birds of prey as a model group, we examine how one of these new morphometric methods – eigensurface analysis – can be used to investigate adaptation. Virtual models of the humerus of 50 falconiform species from 29 genera were examined to identify the morphological correlates associated with different flight styles, habitats, and behaviours of these ecologically sensitive predators. Results indicate that strong, consistent, unique, and subtle modes of shape variation are associated with differences in flight speed (fast–slow), flight style (perch hunting, chasing, low flight, soaring flight, hovering), habitat (forested–open landscape) and migratory behaviour (long-distance migrants–sedentary species). In each case, visual modelling of between-groups transitions in morphology can be used to facilitate identification of features of the humerus, such as the size and extent of surface-shape variation within specific regions of muscle attachment, that are important predictors of function and lifestyle. Eigensurface analysis-based representations of 3D morphology, in combination with standard linear discrimination techniques and new shape modelling procedures, represent a way to statistically evaluate hypotheses of morphological adaptation. This approach can be used not only for bird taxa, but more broadly in studies of ecology and adaptation in many vertebrate, invertebrate, and plant species, in ways that cannot be duplicated either by visual observation or by the representation of form using linear measurements, 2D–3D landmark coordinate sets or sets of 2D–3D boundary outline semilandmarks. © 2012 The Trustees of the Natural History Museum. Zoological Journal of the Linnean Society© 2012 The Linnean Society of London, 165, 390–419.

Aerodynamic flight performance in flap-gliding birds and bats

Many birds use a flight mode called undulating or flap-gliding flight, where they alternate between flapping and gliding phases, while only a few bats make use of such a flight mode. Among birds, flap-gliding is commonly used by medium to large species, where it is regarded to have a lower energetic cost than continuously flapping flight.Here, we introduce a novel model for estimating the energetic flight economy of flap-gliding animals, by determining the lift-to-drag ratio for flap-gliding based on empirical lift-to-drag ratio estimates for continuous flapping flight and for continuous gliding flight, respectively. We apply the model to flight performance data of the common swift (Apus apus) and of the lesser long-nosed bat (Leptonycteris yerbabuenae). The common swift is a typical flap-glider while—to the best of our knowledge—the lesser long-nosed bat does not use flap-gliding.The results show that, according to the model, the flap-gliding common swift saves up to 15% energy compared to a continuous flapping swift, and that this is primarily due to the exceptionally high lift-to-drag ratio in gliding flight relative to that in flapping flight for common swifts. The lesser long-nosed bat, on the other hand, seems not to be able to reduce energetic costs by flap-gliding. The difference in relative costs of flap-gliding flight between the common swift and the lesser long-nosed bat can be explained by differences in morphology, flight style and wake dynamics.The model presented here proves to be a valuable tool for estimating energetic flight economy in flap-gliding animals. The results show that flap-gliding flight that is naturally used by common swifts is indeed the most economic one of the two flight modes, while this is not the case for the non-flap-gliding lesser long-nosed bat.

Size scaling and stiffness of avian primary feathers: implications for the flight of Mesozoic birds

The primary feathers of birds are subject to cyclical forces in flight causing their shafts (rachises) to bend. The amount the feathers deflect during flight is dependent upon the flexural stiffness of the rachises. By quantifying scaling relationships between body mass and feather linear dimensions in a large data set of living birds, we show that both feather length and feather diameter scale much closer to predictions for geometric similarity than they do to elastic similarity. Scaling allometry also indicates that the primary feathers of larger birds are relatively shorter and their rachises relatively narrower, compared to those of smaller birds. Two-point bending tests indicated that larger birds have more flexible feathers than smaller species. Discriminant functional analyses (DFA) showed that body mass, primary feather length and rachis diameter can be used to differentiate between different magnitudes of feather bending stiffness, with primary feather length explaining 63% of variance in rachis stiffness. Adding fossil measurement data to our DFA showed that Archaeopteryx and Confuciusornis do not overlap with extant birds. This strongly suggests that the bending stiffness of their primary feathers was different to extant birds and provides further evidence for distinctive flight styles and likely limited flight ability in Archaeopteryx and Confuciusornis.

Flexural stiffness of feather shafts: geometry rules over material properties

Flight feathers of birds interact with the flow field during flight. They bend and twist under aerodynamic loads. Two parameters are mainly responsible for flexibility in feathers: the elastic modulus (Young's modulus, E) of the material (keratin) and the geometry of the rachises, more precisely the second moment of area (I). Two independent methods were employed to determine Young's modulus of feather rachis keratin. Moreover, the second moment of area and the bending stiffness of feather shafts from fifth primaries of barn owls (Tyto alba) and pigeons (Columba livia) were calculated. These species of birds are of comparable body mass but differ in wing size and flight style. Whether their feather material (keratin) underwent an adaptation in stiffness was previously unknown. This study shows that no significant variation in Young's modulus between the two species exists. However, differences in Young's modulus between proximal and distal feather regions were found in both species. Cross-sections of pigeon rachises were particularly well developed and rich in structural elements, exemplified by dorsal ridges and a well-pronounced transversal septum. In contrast, cross-sections of barn owl rachises were less profiled but had a higher second moment of area. Consequently, the calculated bending stiffness (EI) was higher in barn owls as well. The results show that flexural stiffness is predominantly influenced by the geometry of the feathers rather than by local material properties.

Species-Specific Flight Styles of Flies are Reflected in the Response Dynamics of a Homolog Motion-Sensitive Neuron.

Hoverflies and blowflies have distinctly different flight styles. Yet, both species have been shown to structure their flight behavior in a way that facilitates extraction of 3D information from the image flow on the retina (optic flow). Neuronal candidates to analyze the optic flow are the tangential cells in the third optical ganglion – the lobula complex. These neurons are directionally selective and integrate the optic flow over large parts of the visual field. Homolog tangential cells in hoverflies and blowflies have a similar morphology. Because blowflies and hoverflies have similar neuronal layout but distinctly different flight behaviors, they are an ideal substrate to pinpoint potential neuronal adaptations to the different flight styles. In this article we describe the relationship between locomotion behavior and motion vision on three different levels: (1) We compare the different flight styles based on the categorization of flight behavior into prototypical movements. (2) We measure the species-specific dynamics of the optic flow under naturalistic flight conditions. We found the translational optic flow of both species to be very different. (3) We describe possible adaptations of a homolog motion-sensitive neuron. We stimulate this cell in blowflies (Calliphora) and hoverflies (Eristalis) with naturalistic optic flow generated by both species during free flight. The characterized hoverfly tangential cell responds faster to transient changes in the optic flow than its blowfly homolog. It is discussed whether and how the different dynamical response properties aid optic flow analysis.

Avian wing proportions and flight styles: first step towards predicting the flight modes of mesozoic birds.

We investigated the relationship between wing element proportions and flight mode in a dataset of living avian species to provide a framework for making basic estimates of the range of flight styles evolved by Mesozoic birds. Our results show that feather length (f prim) and total arm length (ta) (sum of the humerus, ulna and manus length) ratios differ significantly between four flight style groups defined and widely used for living birds and as a result are predictive for fossils. This was confirmed using multivariate ordination analyses, with four wing elements (humerus, ulna/radius, manus, primary feathers), that discriminate the four broad flight styles within living birds. Among the variables tested, manus length is closely correlated with wing size, yet is the poorest predictor for flight style, suggesting that the shape of the bones in the hand wing is most important in determining flight style. Wing bone thickness (shape) must vary with wing beat strength, with weaker forces requiring less bone. Finally, we show that by incorporating data from Mesozoic birds, multivariate ordination analyses can be used to predict the flight styles of fossils.

Energy expenditures for flight, aerodynamic quality, and colonization of forest habitats by birds

The power that the birds can use for flight (available power) and the power required for flight according to physical laws (requisite power) grow with an increase in body mass, the exponents of the corresponding functions being different. Small birds can follow different strategies, either improving the aerodynamic quality of the body (thereby saving the excess available power) or sacrifice aerodynamic quality in favor of morphological adaptation to factors other than the demands of flight proper, which provides the possibility of utilizing a wider range of ecological niches. A hypothesis is proposed that the high metabolic rate of passerine birds, compared to representatives of other bird orders, is an adaptation to maneuverable (i.e., relatively low-speed) flight necessary for successful colonization of forest habitats. The speed that birds of such size can develop according to the scaling theory is too high for nesting and foraging in tree crowns, and its reduction is possible in two ways: by increasing air drag or by changing the style of flight (by analogy with airplane vs. helicopter). The first way is feasible, but a high air drag due to morphological modifications (e.g., in the size of the tail or characteristics of the wing) interferes with the possibility of long-distance migration flight, as energy expenditures for it will exceed the energy potential of the bird. This is why migratory nonpasserine birds, which have used this strategy, are practically absent in forests of the temperate zone. Therefore, more promising is the second way involving transition to a new flight style and, in a certain sense, to a new morphophysiological organization. Passerines have achieved this by changing their flight style so that the wing actively generates forces (lift and thrust) only in downstroke. Such a flight requires more energy, and, to provide it in sufficient amounts, passerine birds have increased their basal metabolic rate (BMR). Thus, both their flight energy expenditures and BMR are higher than in nonpasserines. Remarkably, among approximately 8660 extant bird species known today, more than half (about 5100 species) belong to the order Passeriformes. Such a ratio, unknown in any other vertebrate class, is evidence that passerines have gained a considerable biological advantage over all other birds due to their increased BMR.

A comparative study of the mechanics of the pectoralis muscle of the red‐tailed hawk and the barred owl

A comparison of the isometric forces and levers of the pectoralis muscle in red-tailed hawks (Buteo jamaicensis) and barred owls (Strix varia) was done to identify differences that may correlate with their different flight styles. The pectoralis consists of two heads, the anterior m. sternobrachialis (SB) and the posterior m. thoracobrachialis (TB). These are joined at an intramuscular tendon and are supplied by separate primary nerve branches. As in other birds, the two heads have distinct fiber orientations in red-tailed hawks and barred owls. SB's fiber orientation (posterolateral and mediolateral from origin to insertion) provides pronation and protraction of the humerus during adduction. Electromyographic studies in pigeons show that it is active in early downstroke and during level flight. TB is more active during take-off and landing in pigeons. The anterolateral orientation (from origin to insertion) of its fibers provides a retractive component to humeral adduction used to control the wing during landing. In our study, the maximum isometric force produced by the combined pectoralis heads did not differ significantly between the hawk and owl, however, the forces were distributed differently between the two muscle heads. In the owl, SB and TB were capable of producing equal amounts of force, but in the hawk, SB produced significantly less force than did TB. This may reflect the need for a large TB to control landing in both birds during prey-strike, with the owl maintaining both protractive (using SB) and retractive (using TB) abilities. Pronation and protraction may be less important in the flight behavior of the hawk, but its prey-strike behavior may require the maintenance of a substantial TB for braking and controlled stalling, as it initiates strike behavior.

Scaling of avian primary feather length.

The evolution of the avian wing has long fascinated biologists, yet almost no work includes the length of primary feathers in consideration of overall wing length variation. Here we show that the length of the longest primary feather () contributing to overall wing length scales with negative allometry against total arm (ta = humerus+ulna+manus). The scaling exponent varied slightly, although not significantly so, depending on whether a species level analysis was used or phylogeny was controlled for using independent contrasts: . The scaling exponent was not significantly different from that predicted (0.86) by earlier work. It appears that there is a general trend for the primary feathers of birds to contribute proportionally less, and ta proportionally more, to overall wingspan as this dimension increases. Wingspan in birds is constrained close to mass (M 1/3) because of optimisation for lift production, which limits opportunities for exterior morphological change. Within the wing, variations in underlying bone and feather lengths nevertheless may, in altering the joint positions, permit a range of different flight styles by facilitating variation in upstroke kinematics.

An association between ear and tail morphologies of bats and their foraging style

Most studies relating bat morphology to flight ecology have concentrated on the wing membrane. Here, canonical variance analysis showed that the ear and tail morphologies of bats also strongly relate to foraging strategy, which in turn is correlated with flight style. Variations in tail membrane morphology are likely to be a trade-off between increases in the mechanical cost of flight and improvements in foraging and flight performance. Flying with large ears is also potentially energetically expensive, particularly at high flight speeds. Large ears, therefore, are only likely to be affordable for slow foraging gleaning bat species. Bats with faster foraging flight styles tend to have smaller ears, possibly to cut the overall drag produced and reduce the power required for flight. Variations in the size of ears and tail membranes appear to be driven primarily by foraging strategy and not by body size, because the scaling relationships found are either weak or not significant. Ear size in bats may be a result of a trade-off between acoustic and aerodynamic performance.

A syntax of hoverfly flight prototypes

Hoverflies such as Eristalis tenax Linnaeus are known for their distinctive flight style. They can hover on the same spot for several seconds and then burst into movement in apparently any possible direction. In order to determine a quantitative and structured description of complex flight manoeuvres, we searched for a set of repeatedly occurring prototypical movements (PMs) and a set of rules for their ordering. PMs were identified by applying clustering algorithms to the translational and rotational velocities of the body of Eristalis during free-flight sequences. This approach led to nine stable and reliable PMs, and thus provided a tremendous reduction in the complexity of behavioural description. This set of PMs together with the probabilities of transition between them constitute a syntactical description of flight behaviour. The PMs themselves can be roughly segregated into fast rotational turns (saccades) and a variety of distinct translational movements (intersaccadic intervals). We interpret this segregation as reflecting an active sensing strategy which facilitates the extraction of spatial information from retinal image displacements. Detailed analysis of saccades shows that they are performed around all rotational axes individually and in all possible combinations. We found the probability of occurrence of a given saccade type to depend on parameters such as the angle between the long body axis and the direction of flight.

Scaling of Soaring Seabirds and Implications for Flight Abilities of Giant Pterosaurs

The flight ability of animals is restricted by the scaling effects imposed by physical and physiological factors. In comparisons of the power available from muscle and the mechanical power required to fly, it is predicted that the margin between the powers should decrease with body size and that flying animals have a maximum body size. However, predicting the absolute value of this upper limit has proven difficult because wing morphology and flight styles varies among species. Albatrosses and petrels have long, narrow, aerodynamically efficient wings and are considered soaring birds. Here, using animal-borne accelerometers, we show that soaring seabirds have two modes of flapping frequencies under natural conditions: vigorous flapping during takeoff and sporadic flapping during cruising flight. In these species, high and low flapping frequencies were found to scale with body mass (mass −0.30 and mass −0.18) in a manner similar to the predictions from biomechanical flight models (mass −1/3 and mass −1/6). These scaling relationships predicted that the maximum limits on the body size of soaring animals are a body mass of 41 kg and a wingspan of 5.1 m. Albatross-like animals larger than the limit will not be able to flap fast enough to stay aloft under unfavourable wind conditions. Our result therefore casts doubt on the flying ability of large, extinct pterosaurs. The largest extant soarer, the wandering albatross, weighs about 12 kg, which might be a pragmatic limit to maintain a safety margin for sustainable flight and to survive in a variable environment.

Aerodynamics of intermittent bounds in flying birds

Flap-bounding is a common flight style in small birds in which flapping phases alternate with flexed-wing bounds. Body lift is predicted to be essential to making this flight style an aerodynamically attractive flight strategy. To elucidate the contributions of the body and tail to lift and drag during the flexed-wing bound phase, we used particle image velocimetry (PIV) and measured properties of the wake of zebra finch (Taeniopygia guttata, N = 5), flying at 6-10 m s− 1 in a variable speed wind tunnel as well as flow around taxidermically prepared specimens (N = 4) mounted on a sting instrumented with force transducers. For the specimens, we varied air velocity from 2 to 12 m s− 1 and body angle from –15∘ to 50∘. The wake of bounding birds and mounted specimens consisted of a pair of counterrotating vortices shed into the wake from the tail, with induced downwash in the sagittal plane and upwash in parasagittal planes lateral to the bird. This wake structure was present even when the tail was entirely removed. We observed good agreement between force measures derived from PIV and force transducers over the range of body angles typically used by zebra finch during forward flight. Body lift:drag (L:D) ratios averaged 1.4 in live birds and varied between 1 and 1.5 in specimens at body angles from 10∘ to 30∘. Peak (L:D) ratio was the same in live birds and specimens (1.5) and was exhibited in specimens at body angles of 15∘ or 20∘, consistent with the lower end of body angles utilized during bounds. Increasing flight velocity in live birds caused a decrease in C L and C D from maximum values of 1.19 and 0.95 during flight at 6 m s− 1 to minimum values of 0.70 and 0.54 during flight at 10 m s− 1. Consistent with delta-wing theory as applied to birds with a graduated-tail shape, trimming the tail to 0 and 50% of normal length reduced L:D ratios and extending tail length to 150% of normal increased L:D ratio. As downward induced velocity is present in the sagittal plane during upstroke of flapping flight, we hypothesize that body lift is produced during flapping phases. Future efforts to model the mechanics of intermittent flight should take into account that flap-bounding birds may support up to 20% of their weight even with their wings fully flexed.

Scaling of soaring seabirds and its implication for the maximum size of flying pterosaurs

AbstractThe flight ability of animals is restricted by the scaling effects imposed by physical and physiological factors. In comparisons of the power available from muscle and the mechanical power required to fly, theoretical studies have predicted that the margin between the powers should decrease with body size and that flying animals have a maximum body size. However, predicting an absolute value of this upper limit has been difficult because wing morphology and flight styles vary among species. Albatrosses and petrels have long, narrow, aerodynamically efficient wings and are considered to be soaring birds. Here, using animal-borne accelerometers, we show that scaling analyses of wing-flapping frequencies in these seabirds indicate that the maximum size limit for soaring animals is a body mass of 41 kg and a wingspan of 5.1 m. Soaring seabirds were observed to have two modes of flapping frequencies: vigorous flapping during takeoff and sporadic flapping during cruising flight. In these species, high and low flapping frequencies were found to scale with body mass (mass ^-0.30^ and mass ^-0.18^) in a manner similar to the predictions from biomechanical flight models (mass ^-1/3^ and mass ^-1/6^). The scaling relationships predicted that animals larger than the limit will not be able to flap fast enough to stay aloft under unfavourable wind conditions. Our result therefore casts doubt on the flying ability of large, extinct pterosaurs. The largest extant soarer, the wandering albatross, weighs about 10 kg, which might be a pragmatic limit to maintain a safety margin for sustainable flight and to survive in a variable environment.

The fossil record and limb disparity of enantiornithines, the dominant flying birds of the Cretaceous

Morphometric and stratigraphic analyses that encompass the known fossil record of enantiornithine birds (Enantiornithes) are presented. These predominantly flighted taxa were the dominant birds of the second half of the Mesozoic; the enantiornithine lineage is known to have lasted for at least 60 million years (Ma), up until the end of the Cretaceous. Analyses of fossil record dynamics show that enantiornithine ‘collectorship’ since the 1980s approaches an exponential distribution, indicating that an asymptote in proportion of specimens has yet to be achieved. Data demonstrate that the fossil record of enantiornithines is complete enough for the extraction of biological patterns. Comparison of the available fossil specimens with a large data set of modern bird (Neornithes) limb proportions also illustrates that the known forelimb proportions of enantiornithines fall within the range of extant taxa; thus these birds likely encompassed the range of flight styles of extant birds. In contrast, most enantiornithines had hindlimb proportions that differ from any extant taxa. To explore this, ternary diagrams are used to graph enantiornithine limb variation and to identify some morphological oddities (Otogornis, Gobipteryx); taxa not directly comparable to modern birds. These exceptions are interesting – although anatomically uniform, and similar to extant avians in their wing proportions, some fossil enantiornithines likely had flight styles not seen among their living counterparts.

Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation

Bats (Chiroptera) represent one of the largest and most diverse radiations of mammals, accounting for one-fifth of extant species. Although recent studies unambiguously support bat monophyly and consensus is rapidly emerging about evolutionary relationships among extant lineages, the fossil record of bats extends over 50 million years, and early evolution of the group remains poorly understood. Here we describe a new bat from the Early Eocene Green River Formation of Wyoming, USA, with features that are more primitive than seen in any previously known bat. The evolutionary pathways that led to flapping flight and echolocation in bats have been in dispute, and until now fossils have been of limited use in documenting transitions involved in this marked change in lifestyle. Phylogenetically informed comparisons of the new taxon with other bats and non-flying mammals reveal that critical morphological and functional changes evolved incrementally. Forelimb anatomy indicates that the new bat was capable of powered flight like other Eocene bats, but ear morphology suggests that it lacked their echolocation abilities, supporting a 'flight first' hypothesis for chiropteran evolution. The shape of the wings suggests that an undulating gliding-fluttering flight style may be primitive for bats, and the presence of a long calcar indicates that a broad tail membrane evolved early in Chiroptera, probably functioning as an additional airfoil rather than as a prey-capture device. Limb proportions and retention of claws on all digits indicate that the new bat may have been an agile climber that employed quadrupedal locomotion and under-branch hanging behaviour.