Vertebrate Flight Research Articles

Bats use topography and nocturnal updrafts to fly high and fast

During the day, flying animals exploit the environmental energy landscape by seeking out thermal or orographic uplift, or extracting energy from wind gradients.1-6 However, most of these energy sources are not thought to be available at night because of the lower thermal potential in the nocturnal atmosphere, as well as the difficulty of locating features that generate uplift. Despite this, several bat species have been observed hundreds to thousands of meters above the ground.7-9 Individuals make repeated, energetically costly high-altitude ascents,10-13 and others fly at some of the fastest speeds observed for powered vertebrate flight.14 We hypothesized that bats use orographic uplift to reach high altitudes,9,15-17 and that both this uplift and bat high-altitude ascents would be highly predictable.18 By superimposing detailed three-dimensional GPS tracking of European free-tailed bats (Tadarida teniotis) on high-resolution regional wind data, we show that bats do indeed use the energy of orographic uplift to climb to over 1,600 m, and also that they reach maximum sustained self-powered airspeeds of 135km h-1. We show that wind and topography can predict areas of the landscape able to support high-altitude ascents, and that bats use these locations to reach high altitudes while reducing airspeeds. Bats then integrate wind conditions to guide high-altitude ascents, deftly exploiting vertical wind energy in the nocturnal landscape.

Evolution of Flight Muscle Contractility and Energetic Efficiency.

The powered flight of animals requires efficient and sustainable contractions of the wing muscles of various flying species. Despite their high degree of phylogenetic divergence, flight muscles in insects and vertebrates are striated muscles with similarly specialized sarcomeric structure and basic mechanisms of contraction and relaxation. Comparative studies examining flight muscles together with other striated muscles can provide valuable insights into the fundamental mechanisms of muscle contraction and energetic efficiency. Here, we conducted a literature review and data mining to investigate the independent emergence and evolution of flight muscles in insects, birds, and bats, and the likely molecular basis of their contractile features and energetic efficiency. Bird and bat flight muscles have different metabolic rates that reflect differences in energetic efficiencies while having similar contractile machinery that is under the selection of similar natural environments. The significantly lower efficiency of insect flight muscles along with minimized energy expenditure in Ca2+ handling is discussed as a potential mechanism to increase the efficiency of mammalian striated muscles. A better understanding of the molecular evolution of myofilament proteins in the context of physiological functions of invertebrate and vertebrate flight muscles can help explore novel approaches to enhance the performance and efficiency of skeletal and cardiac muscles for the improvement of human health.

Speciation Associated with Shifts in Migratory Behavior in an Avian Radiation.

Migratory animals move up to thousands of kilometers every year [1]. Losses of migration (i.e., migratory drop-offs) occur when individuals of a migratory species stop migrating and establish founder sedentary populations, a phenomenon documented in birds [2-5] and butterflies [6]. In theory, losses-and also gains-of migration might promote speciation if sedentary and migratory populations become reproductively isolated [7-9]. Because migratory and sedentary strategies involve alternative physiological, behavioral, and morphological traits [10-13], divergence along multiple axes of organismal function is expected to accompany switches in migratory behavior, potentially accelerating speciation. We present evidence of speciation driven by a migratory drop-off in the fork-tailed flycatcher (Tyrannus savana) resulting in reproductive isolation likely driven by changes in breeding schedules (allochronic speciation [13-15]) and geographic isolation of breeding grounds (allopatric speciation [16]). Phylogenetic analyses across New World flycatchers (Tyrannidae) showed that an association between speciation and drop-offs is also observable at a macroevolutionary scale. Loss of migration was significantly more frequent than its gain, and speciation rates of migratory and partially migratory lineages (i.e., species having both migratory and sedentary populations) exceeded those of sedentary lineages. Models of trait evolution indicated that partial migration is an intermediate step between migratory and sedentary states in this family. Given that partial migration is widespread across migratory animals (e.g., of all migratory birds, ca. 51% are partially migratory [5]), speciation via switches in migratory behavior might be an important yet overlooked mechanism of animal diversification.

Postcranial heterochrony, modularity, integration and disparity in the prenatal ossification in bats (Chiroptera)

BackgroundSelf-powered flight is one of the most energy-intensive types of locomotion found in vertebrates. It is also associated with a range of extreme morpho-physiological adaptations that evolved independently in three different vertebrate groups. Considering that development acts as a bridge between the genotype and phenotype on which selection acts, studying the ossification of the postcranium can potentially illuminate our understanding of bat flight evolution. However, the ontogenetic basis of vertebrate flight remains largely understudied. Advances in quantitative analysis of sequence heterochrony and morphogenetic growth have created novel approaches to study the developmental basis of diversification and the evolvability of skeletal morphogenesis.Assessing the presence of ontogenetic disparity, integration and modularity from an evolutionary approach allows assessing whether flight may have resulted in evolutionary differences in the magnitude and mode of development in bats.ResultsWe quantitatively compared the prenatal ossification of the postcranium (24 bones) between bats (14 species), non-volant mammals (11 species) and birds (14 species), combining for the first time prenatal sequence heterochrony and developmental growth data. Sequence heterochrony was found across groups, showing that bat postcranial development shares patterns found in other flying vertebrates but also those in non-volant mammals. In bats, modularity was found as an axial-appendicular partition, resembling a mammalian pattern of developmental modularity and suggesting flight did not repattern prenatal postcranial covariance in bats.ConclusionsCombining prenatal data from 14 bat species, this study represents the most comprehensive quantitative analysis of chiropteran ossification to date. Heterochrony between the wing and leg in bats could reflect functional needs of the newborn, rather than ecological aspects of the adult. Bats share similarities with birds in the development of structures involved in flight (i.e. handwing and sternum), suggesting that flight altriciality and early ossification of pedal phalanges and sternum are common across flying vertebrates. These results indicate that the developmental modularity found in bats facilitates intramodular phenotypic diversification of the skeleton. Integration and disparity increased across developmental time in bats. We also found a delay in the ossification of highly adaptable and evolvable regions (e.g. handwing and sternum) that are directly associated with flight performance.

Inspiration for wing design: how forelimb specialization enables active flight in modern vertebrates.

Harnessing flight strategies refined by millions of years of evolution can help expedite the design of more efficient, manoeuvrable and robust flying robots. This review synthesizes recent advances and highlights remaining gaps in our understanding of how bird and bat wing adaptations enable effective flight. Included in this discussion is an evaluation of how current robotic analogues measure up to their biological sources of inspiration. Studies of vertebrate wings have revealed skeletal systems well suited for enduring the loads required during flight, but the mechanisms that drive coordinated motions between bones and connected integuments remain ill-described. Similarly, vertebrate flight muscles have adapted to sustain increased wing loading, but a lack of in vivo studies limits our understanding of specific muscular functions. Forelimb adaptations diverge at the integument level, but both bird feathers and bat membranes yield aerodynamic surfaces with a level of robustness unparalleled by engineered wings. These morphological adaptations enable a diverse range of kinematics tuned for different flight speeds and manoeuvres. By integrating vertebrate flight specializations-particularly those that enable greater robustness and adaptability-into the design and control of robotic wings, engineers can begin narrowing the wide margin that currently exists between flying robots and vertebrates. In turn, these robotic wings can help biologists create experiments that would be impossible in vivo.

Flapping wing aerodynamics: from insects to vertebrates.

More than a million insects and approximately 11,000 vertebrates utilize flapping wings to fly. However, flapping flight has only been studied in a few of these species, so many challenges remain in understanding this form of locomotion. Five key aerodynamic mechanisms have been identified for insect flight. Among these is the leading edge vortex, which is a convergent solution to avoid stall for insects, bats and birds. The roles of the other mechanisms - added mass, clap and fling, rotational circulation and wing-wake interactions - have not yet been thoroughly studied in the context of vertebrate flight. Further challenges to understanding bat and bird flight are posed by the complex, dynamic wing morphologies of these species and the more turbulent airflow generated by their wings compared with that observed during insect flight. Nevertheless, three dimensionless numbers that combine key flow, morphological and kinematic parameters - the Reynolds number, Rossby number and advance ratio - govern flapping wing aerodynamics for both insects and vertebrates. These numbers can thus be used to organize an integrative framework for studying and comparing animal flapping flight. Here, we provide a roadmap for developing such a framework, highlighting the aerodynamic mechanisms that remain to be quantified and compared across species. Ultimately, incorporating complex flight maneuvers, environmental effects and developmental stages into this framework will also be essential to advancing our understanding of the biomechanics, movement ecology and evolution of animal flight.

Challenges and advances in the study of pterosaur flight

Pterosaurs have fascinated scientists and nonscientists alike for over 200 years, as one of the three known clades of vertebrates to have evolved flapping flight. The smallest pterosaurs were comparable in size to the smallest extant birds and bats, but the largest pterosaurs were vastly larger than any extant flier. This immense size range, coupled with poor preservation and adaptations for flight unknown in extant vertebrates, have made interpretations of pterosaur flight problematic and often contentious. Here we review the anatomical, evolutionary, and phylogenetic history of pterosaurs, as well as the views, perspectives, and biases regarding their interpretation. In recent years, three areas of pterosaur biology have faced challenges and made advances: structure of the wing membrane, function of the pteroid, body size and mass estimates, as well as flight mechanics and aerodynamics. Comparative anatomical and fossil study, simulated bone loading, and aerodynamic modeling have all proved successful in furthering our understanding of pterosaur flight. We agree with previous authors that pterosaurs should be studied as pterosaurs, a diverse but phylogenetically, anatomically, and mechanically constrained clade that can offer new insights into the diversity of vertebrate flight.

Power of the wingbeat: modelling the effects of flapping wings in vertebrate flight.

Animal flight performance has been studied using models developed for man-made aircraft. For an aeroplane with fixed wings, the energetic cost as a function of flight speed can be expressed in terms of weight, wing span, wing area and body area, where more details are included in proportionality coefficients. Flying animals flap their wings to produce thrust. Adopting the fixed wing flight model implicitly incorporates the effects of wing flapping in the coefficients. However, in practice, these effects have been ignored. In this paper, the effects of reciprocating wing motion on the coefficients of the fixed wing aerodynamic power model for forward flight are explicitly formulated in terms of thrust requirement, wingbeat frequency and stroke-plane angle, for optimized wingbeat amplitudes. The expressions are obtained by simulating flights over a large parameter range using an optimal vortex wake method combined with a low-level blade element method. The results imply that previously assumed acceptable values for the induced power factor might be strongly underestimated. The results also show the dependence of profile power on wing kinematics. The expressions introduced in this paper can be used to significantly improve animal flight models.

Wing bone laminarity is not an adaptation for torsional resistance in bats.

Torsional loading is a common feature of skeletal biomechanics during vertebrate flight. The importance of resisting torsional loads is best illustrated by the convergence of wing bone structure (e.g., long with thin walls) across extant bats and birds. Whether or not such a convergence occurs at the microstructural level is less clear. In volant birds, the humeri and ulnae often contain abundant laminar bony tissue in which primary circumferential vascular canals course concentrically about the long axis of the bone. These circumferential canals and the matrix surrounding them presumably function to resist the tissue-level shear stress caused by flight-induced torsion. Here, we assess whether or not laminar bone is a general adaptive feature in extant flying vertebrates using a histological analysis of bat bones. We sampled the humeri from six adult taxa representing a broad phylogenetic and body size range (6–1,000 g). Transverse thick sections were prepared from the midshaft of each humerus. Bone tissue was classified based on the predominant orientation of primary vascular canals. Our results show that humeri from bats across a wide phylogenetic and body size range do not contain any laminar bone. Instead, humeri are essentially avascular in bats below about 100 g and are poorly vascularized with occasional longitudinal to slightly radial canals in large bats. In contrast, humeri from birds across a comparable size range (40–1,000 g) are highly vascularized with a wide range in bone laminarity. Phylogenetically-informed scaling analyses reveal that the difference in vascularity between birds and bats is best explained by higher somatic relative growth rates in birds. The presence of wing bone laminarity in birds and its absence in bats suggests that laminar bone is not a necessary biomechanical feature in flying vertebrates and may be apomorphic to birds.

A sensory-motor control model of animal flight explains why bats fly differently in light versus dark.

Animal flight requires fine motor control. However, it is unknown how flying animals rapidly transform noisy sensory information into adequate motor commands. Here we developed a sensorimotor control model that explains vertebrate flight guidance with high fidelity. This simple model accurately reconstructed complex trajectories of bats flying in the dark. The model implies that in order to apply appropriate motor commands, bats have to estimate not only the angle-to-target, as was previously assumed, but also the angular velocity (“proportional-derivative” controller). Next, we conducted experiments in which bats flew in light conditions. When using vision, bats altered their movements, reducing the flight curvature. This change was explained by the model via reduction in sensory noise under vision versus pure echolocation. These results imply a surprising link between sensory noise and movement dynamics. We propose that this sensory-motor link is fundamental to motion control in rapidly moving animals under different sensory conditions, on land, sea, or air.

Straight-line climbing flight aerodynamics of a fruit bat

From flight data obtained on a fruit bat, Cynopterus brachyotis, a kinematic model for straight-line flapping motion is extracted and analyzed in a computational fluid dynamics (CFD) framework to gain insight into the complexity of bat flight. The intricate functional mechanics and architecture of the bat wings set it apart from other vertebrate flight. The extracted kinematic model is simulated for a range of Reynolds numbers, to observe the effect these phenomena have on the unsteady transient mechanisms of the flow produced by the flapping wings. The Strouhal number calculated from the data is high indicating that the oscillatory motion dominates the flow physics. From the obtained data, the bat exhibits fine control of its mechanics by actively varying wing camber, wing area, torsional rotation of the wing, forward and backward translational sweep of the wing, and wing conformation to dictate the fluid dynamics. As is common in flapping flight, the primary force generation is through the attached unsteady vortices on the wing surface. The bat through varying the wing camber and the wing area modulates this force output. The power requirement for the kinematics is analyzed and correlated with the aerodynamic performance.

An aeroelastic instability provides a possible basis for the transition from gliding to flapping flight.

The morphology, kinematics and stiffness properties of lifting surfaces play a key role in the aerodynamic performance of vertebrate flight. These surfaces, as a result of their flexible nature, may move both actively, owing to muscle contraction, and passively, in reaction to fluid forces. However, the nature and implications of this fluid-structure interaction are not well understood. Here, we study passive flight (flight with no active wing actuation) and explore a physical mechanism that leads to the emergence of a natural flapping motion. We model a vertebrate wing with a compliant shoulder and the ability to camber with an idealized physical model consisting of a cantilevered flat plate with a hinged trailing flap. We find that at low wind speed the wing is stationary, but at a critical speed the wing spontaneously flaps. The lift coefficient is significantly enhanced once the wing starts to oscillate, although this increase in lift generation is accompanied by an increase in drag. Flow visualization suggests that a strong leading edge vortex attached to the wing during downstroke is the primary mechanism responsible for the enhanced lift. The flapping instability we observe suggests a possible scenario for an evolutionary transition from gliding to powered flapping flight in animals that possess compliant wings capable of passive camber. Although the flapping state is accompanied by a lower lift-to-drag ratio, the increased lifting capability it confers might have enabled increased body mass, improved foraging performance and/or flight at lower speeds, any of which might have been selectively advantageous.

Move That Fatty Acid: Fuel Selection and Transport in Migratory Birds and Bats

The metaphor of marathon running is inadequate to fully capture the magnitude of long-distance migratory flight of birds. In some respects a journey to the moon seems more appropriate. Birds have no access to supplementary water or nutrition during a multi-day flight, and they must carefully budget their body fat and protein stores to provide both fuel and life support. Fatty acid transport is crucial to successful non-stop migratory flight in birds. Although fat is the most energy-dense metabolic fuel, the insolubility of its component fatty acids makes them difficult to transport to working muscles fast enough to support the highly aerobic exercise required to fly. Recent evidence indicates that migratory birds compensate for this by expressing large amounts of fatty acid transport proteins on the membranes of the muscles (FAT/CD36 and FABPpm) and in the cytosol (H-FABP). Through endogenous mechanisms and/or diet, migratory birds may alter the fatty acid composition of the fat stores and muscle membranes to improve endurance during flight. Fatty acid chain length, degree of unsaturation, and placement of double bonds can affect the rate of mobilization of fatty acids from adipose tissue, utilization of fatty acids by muscles, and whole-animal performance. However, there is great uncertainty about how important fatty acid composition is to the success of migration or whether particular types of fatty acids (e.g., omega-3 or omega-6) are most beneficial. Migratory bats provide an interesting example of evolutionary convergence with birds, which may provide evidence for the generality of the bird model to the evolution of migration by flight in vertebrates. Yet only recently have attempts been made to study bat migration physiology. Many aspects of their fuel metabolism are predicted to be more similar to those of migrant birds than to those of non-flying mammals. Bats may be distinct from most birds in their potential to conserve energy by using torpor between flights, and in the behavioral and physiological trade-offs they may make between migration and reproduction, which often overlap.

A quantitative comparison of bird and bat wakes

Qualitative comparison of bird and bat wakes has demonstrated significant differences in the structure of the far wake. Birds have been found to have a unified vortex wake of the two wings, while bats have a more complex wake with gradients in the circulation along the wingspan, and with each wing generating its own vortex structure. Here, we compare quantitative measures of the circulation in the far wake of three bird and one bat species. We find that bats have a significantly stronger normalized circulation of the start vortex than birds. We also find differences in how the circulation develops during the wingbeat as demonstrated by the ratio of the circulation of the dominant start vortex and the total circulation of the same sense. Birds show a more prominent change with changing flight speed and a relatively weaker start vortex at minimum power speed than bats. We also find that bats have a higher normalized wake loading based on the start vortex, indicating higher relative induced drag and therefore less efficient lift generation than birds. Our results thus indicate fundamental differences in the aerodynamics of bird and bat flight that will further our understanding of the evolution of vertebrate flight.

The Evolution of Flight in Bats: Narrowing the Field of Plausible Hypotheses

The evolution of flapping flight in bats from an arboreal gliding ancestor appears on the surface to be a relatively simple transition. However, bat flight is a highly complex functional system from a morphological, physiological, and aerodynamic perspective, and the transition from a gliding precursor may involve functional discontinuities that represent evolutionary hurdles. In this review, I suggest a framework for a comprehensive treatment of the evolution of complex functional systems that emphasizes a mechanistic understanding of the initial state, the final state, and the proposed transitional states. In this case, bats represent the final state and extant mammalian gliders are used as a model for the initial state. To explore possible transitional states, I propose a set of criteria for evaluating hypotheses about the evolution of flight in vertebrates and suggest methods by which we can advance our understanding of the transition from gliding to flapping flight. Although it is impossible ever to know with certainty the sequence of events landing to flapping flight, the field of possibilities can be narrowed to those that maintain the functional continuity of the wing and result in improved aerodynamic performance across this transition. The fundamental differences between gliding and flapping flight should not necessarily be seen as evidence that this transition could not occur; rather, these differences point out compelling aspects of the aerodynamics of animal wings that require further investigation.

The nature of flight

Since the dawn of civilization, humans have envied their feathery companions for their ability to leap from the earth, escape gravity—and all earthly problems—and ascend to the skies. Of course, humans eventually made it into the skies too, albeit after a few millennia and rather inelegantly with the help of technology. Even so, we can now fly around the Earth in a matter of hours or zoom about in a fighter plane at supersonic speeds. Yet, flying has never lost its fascination for humans, and many scientists continue to study how various species have mastered the physical and metabolic challenges of flight. Many of the basic questions regarding aerodynamics and the physical mechanics of flight—including the fundamental one of how the bumblebee manages to stay in the air—have largely been resolved. Now, the spotlight has moved to focus on the underlying metabolic and molecular mechanisms that enable animals to fly. ![][1] The ability to fly appears to have evolved separately at least four times: in birds, bats, insects and pterosaurs. Although pterosaurs are extinct, the other three provide unique opportunities to study the aerodynamic and molecular features of animal flight. The evolution of flight in both vertebrates and invertebrates has led to a wide variety of adaptations to exploit unique physiological advantages and to overcome specific deficiencies. For example, size matters and, owing to various mathematical laws, smaller animals have to flap their wings faster to stay in the air. Furthermore, air resistance is a more significant problem for smaller animals, which means that there is an effective minimum size for flight. Yet other factors mean that flying animals also cannot be too large; oxygen and sugars have to travel further around the body, and the power required for take off is proportionately greater—as can be seen by watching how large … [1]: /embed/graphic-1.gif

Gliding and the Functional Origins of Flight: Biomechanical Novelty or Necessity?

A biomechanically parsimonious hypothesis for the evolution of flapping flight in terrestrial vertebrates suggests progression within an arboreal context from jumping to directed aerial descent, gliding with control via appendicular motions, and ultimately to powered flight. The more than 30 phylogenetically independent lineages of arboreal vertebrate gliders lend strong indirect support to the ecological feasibility of such a trajectory. Insect flight evolution likely followed a similar sequence, but is unresolved paleontologically. Recently described falling behaviors in arboreal ants provide the first evidence demonstrating the biomechanical capacity for directed aerial descent in the complete absence of wings. Intentional control of body trajectories as animals fall from heights (and usually from vegetation) likely characterizes many more taxa than is currently recognized. Understanding the sensory and biomechanical mechanisms used by extant gliding animals to control and orient their descent is central to deciphering pathways involved in flight evolution.

The nature of flight

Since the dawn of civilization, humans have envied their feathery companions for their ability to leap from the earth, escape gravity—and all earthly problems—and ascend to the skies. Of course, humans eventually made it into the skies too, albeit after a few millennia and rather inelegantly with the help of technology. Even so, we can now fly around the Earth in a matter of hours or zoom about in a fighter plane at supersonic speeds. Yet, flying has never lost its fascination for humans, and many scientists continue to study how various species have mastered the physical and metabolic challenges of flight. Many of the basic questions regarding aerodynamics and the physical mechanics of flight—including the fundamental one of how the bumblebee manages to stay in the air—have largely been resolved. Now, the spotlight has moved to focus on the underlying metabolic and molecular mechanisms that enable animals to fly. ![][1] The ability to fly appears to have evolved separately at least four times: in birds, bats, insects and pterosaurs. Although pterosaurs are extinct, the other three provide unique opportunities to study the aerodynamic and molecular features of animal flight. The evolution of flight in both vertebrates and invertebrates has led to a wide variety of adaptations to exploit unique physiological advantages and to overcome specific deficiencies. For example, size matters and, owing to various mathematical laws, smaller animals have to flap their wings faster to stay in the air. Furthermore, air resistance is a more significant problem for smaller animals, which means that there is an effective minimum size for flight. Yet other factors mean that flying animals also cannot be too large; oxygen and sugars have to travel further around the body, and the power required for take off is proportionately greater—as can be seen by watching how large … [1]: /embed/graphic-1.gif

Aerodynamic force generation, performance and control of body orientation during gliding in sugar gliders (Petaurus breviceps)

Gliding has often been discussed in the literature as a possible precursor to powered flight in vertebrates, but few studies exist on the mechanics of gliding in living animals. In this study I analyzed the 3D kinematics of sugar gliders (Petaurus breviceps) during short glides in an enclosed space. Short segments of the glide were captured on video, and the positions of marked anatomical landmarks were used to compute linear distances and angles, as well as whole body velocities and accelerations. From the whole body accelerations I estimated the aerodynamic forces generated by the animals. I computed the correlations between movements of the limbs and body rotations to examine the control of orientation during flight. Finally, I compared these results to those of my earlier study on the similarly sized and distantly related southern flying squirrel (Glaucomys volans). The sugar gliders in this study accelerated downward slightly (1.0+/-0.5 m s(-2)), and also accelerated forward (2.1+/-0.6 m s(-2)) in all but one trial, indicating that the body weight was not fully supported by aerodynamic forces and that some of the lift produced forward acceleration rather than just balancing body weight. The gliders used high angles of attack (44.15+/-3.12 degrees ), far higher than the angles at which airplane wings would stall, yet generated higher lift coefficients (1.48+/-0.18) than would be expected for a stalled wing. Movements of the limbs were strongly correlated with body rotations, suggesting that sugar gliders make extensive use of limb movements to control their orientation during gliding flight. In addition, among individuals, different limb movements were associated with a given body rotation, suggesting that individual variation exists in the control of body rotations. Under similar conditions, flying squirrels generated higher lift coefficients and lower drag coefficients than sugar gliders, yet had only marginally shallower glides. Flying squirrels have a number of morphological specializations not shared by sugar gliders that may help to explain their greater lift generating performance.

The evolutionary physiology of animal flight: paleobiological and present perspectives.

Recent geophysical analyses suggest the presence of a late Paleozoic oxygen pulse beginning in the late Devonian and continuing through to the late Carboniferous. During this period, plant terrestrialization and global carbon deposition resulted in a dramatic increase in atmospheric oxygen levels, ultimately yielding concentrations potentially as high as 35% relative to the contemporary value of 21%. Such hyperoxia of the late Paleozoic atmosphere may have physiologically facilitated the initial evolution of insect flight metabolism. Widespread gigantism in late Paleozoic insects and other arthropods is also consistent with enhanced oxygen flux within diffusion-limited tracheal systems. Because total atmospheric pressure increases with increased oxygen partial pressure, concurrently hyperdense conditions would have augmented aerodynamic force production in early forms of flying insects. By the late Permian, evolution of decompositional microbial and fungal communities, together with disequilibrium in rates of carbon deposition, gradually reduced oxygen concentrations to values possibly as low as 15%. The disappearance of giant insects by the end of the Permian is consistent with extinction of these taxa for reasons of asphyxiation on a geological time scale. As with winged insects, the multiple historical origins of vertebrate flight in the late Jurassic and Cretaceous correlate temporally with periods of elevated atmospheric oxygen. Much discussion of flight performance in Archaeopteryx assumes a contemporary atmospheric composition. Elevated oxygen levels in the mid- to late Mesozoic would, however, have facilitated aerodynamic force production and enhanced muscle power output for ancestral birds, as well as for precursors to bats and pterosaurs.