Archaeopteryx is an iconic fossil taxon with feathered wings from the Late Jurassic of Germany that occupies a crucial position for understanding the early evolution of avian flight. After over 150 years of study, its mosaic anatomy unifying characters of both non-flying dinosaurs and flying birds has remained challenging to interpret in a locomotory context. Here, we compare new data from three Archaeopteryx specimens obtained through phase-contrast synchrotron microtomography to a representative sample of archosaurs employing a diverse array of locomotory strategies. Our analyses reveal that the architecture of Archaeopteryx’s wing bones consistently exhibits a combination of cross-sectional geometric properties uniquely shared with volant birds, particularly those occasionally utilising short-distance flapping. We therefore interpret that Archaeopteryx actively employed wing flapping to take to the air through a more anterodorsally posteroventrally oriented flight stroke than used by modern birds. This unexpected outcome implies that avian powered flight must have originated before the latest Jurassic.
The remarkable maneuverability of flying animals results from precise movements of their highly specialized wings. Bats have evolved an impressive capacity to control their flight, in large part due to their ability to modulate wing shape, area, and angle of attack through many independently controlled joints. Bat wings, however, also contain many bones and relatively large muscles, and thus the ratio of bats' wing mass to their body mass is larger than it is for all other extant flyers. Although the inertia in bat wings would typically be associated with decreased aerial maneuverability, we show that bat maneuvers challenge this notion. We use a model-based tracking algorithm to measure the wing and body kinematics of bats performing complex aerial rotations. Using a minimal model of a bat with only six degrees of kinematic freedom, we show that bats can perform body rolls by selectively retracting one wing during the flapping cycle. We also show that this maneuver does not rely on aerodynamic forces, and furthermore that a fruit fly, with nearly massless wings, would not exhibit this effect. Similar results are shown for a pitching maneuver. Finally, we combine high-resolution kinematics of wing and body movements during landing and falling maneuvers with a 52-degree-of-freedom dynamical model of a bat to show that modulation of wing inertia plays the dominant role in reorienting the bat during landing and falling maneuvers, with minimal contribution from aerodynamic forces. Bats can, therefore, use their wings as multifunctional organs, capable of sophisticated aerodynamic and inertial dynamics not previously observed in other flying animals. This may also have implications for the control of aerial robotic vehicles.
Flying lizards of the genus Draco are renowned for their gliding ability, using an aerofoil formed by winglike patagial membranes and supported by elongated thoracic ribs. It remains unknown, however, how these lizards manoeuvre during flight. Here, I present the results of a study on the aerial behaviour of Dussumier’s Flying Lizard (Draco dussumieri) and show that Draco attaches the forelimbs to the leading edge of the patagium while airborne, forming a hitherto unknown type of composite wing. The attachment of the forelimbs to the patagium suggests that that aerofoil is controlled through movements of the forelimbs. One major advantage for the lizards is that the forelimbs retain their complete range of movement and functionality for climbing and running when not used as a part of the wing. These findings not only shed a new light on the flight of Draco but also have implications for the interpretation of gliding performance in fossil species.
- Philosophical transactions of the Royal Society of London. Series B, Biological sciences
- Published over 2 years ago
This work is a synthesis of our current understanding of the mechanics, aerodynamics and visually mediated control of dragonfly and damselfly flight, with the addition of new experimental and computational data in several key areas. These are: the diversity of dragonfly wing morphologies, the aerodynamics of gliding flight, force generation in flapping flight, aerodynamic efficiency, comparative flight performance and pursuit strategies during predatory and territorial flights. New data are set in context by brief reviews covering anatomy at several scales, insect aerodynamics, neuromechanics and behaviour. We achieve a new perspective by means of a diverse range of techniques, including laser-line mapping of wing topographies, computational fluid dynamics simulations of finely detailed wing geometries, quantitative imaging using particle image velocimetry of on-wing and wake flow patterns, classical aerodynamic theory, photography in the field, infrared motion capture and multi-camera optical tracking of free flight trajectories in laboratory environments. Our comprehensive approach enables a novel synthesis of datasets and subfields that integrates many aspects of flight from the neurobiology of the compound eye, through the aeromechanical interface with the surrounding fluid, to flight performance under cruising and higher-energy behavioural modes.This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’.
Measuring the costs of soaring, gliding and flapping flight in raptors is challenging, but essential for understanding their ecology. Among raptors, vultures are scavengers that have evolved highly efficient soaring-gliding flight techniques to minimize energy costs to find unpredictable food resources. Using electrocardiogram, GPS and accelerometer bio-loggers, we report the heart rate (HR) of captive griffon vultures (Gyps fulvus and G. himalayensis) trained for freely-flying. HR increased three-fold at take-off (characterized by prolonged flapping flight) and landing (>300 beats-per-minute, (bpm)) compared to baseline levels (80-100 bpm). However, within 10 minutes after the initial flapping phase, HR in soaring/gliding flight dropped to values similar to baseline levels, i.e. slightly lower than theoretically expected. However, the extremely rapid decrease in HR was unexpected, when compared with other marine gliders, such as albatrosses. Weather conditions influenced flight performance and HR was noticeably higher during cloudy compared to sunny conditions when prolonged soaring flight is made easier by thermal ascending air currents. Soaring as a cheap locomotory mode is a crucial adaptation for vultures who spend so long on the wing for wide-ranging movements to find food.
External morphology is commonly used to identify bats as well as to investigate flight and foraging behavior, typically relying on simple length and area measures or ratios. However, geometric morphometrics is increasingly used in the biological sciences to analyse variation in shape and discriminate among species and populations. Here we compare the ability of traditional versus geometric morphometric methods in discriminating between closely related bat species - in this case European horseshoe bats (Rhinolophidae, Chiroptera) - based on morphology of the wing, body and tail. In addition to comparing morphometric methods, we used geometric morphometrics to detect interspecies differences as shape changes. Geometric morphometrics yielded improved species discrimination relative to traditional methods. The predicted shape for the variation along the between group principal components revealed that the largest differences between species lay in the extent to which the wing reaches in the direction of the head. This strong trend in interspecific shape variation is associated with size, which we interpret as an evolutionary allometry pattern.
It is essential for birds to be agile and aware of their immediate environment, especially when flying through dense foliage. To investigate the type of visual signals and strategies used by birds while negotiating cluttered environments, we presented budgerigars with vertically oriented apertures of different widths. We find that, when flying through narrow apertures, birds execute their maneuvers in an anticipatory fashion, with wing closures, if necessary, occurring well in advance of the aperture. When passing through an aperture that is narrower than the wingspan, the birds close their wings at a specific, constant distance before the aperture, which is independent of aperture width. In these cases, the birds also fly significantly higher, possibly pre-compensating for the drop in altitude. The speed of approach is largely constant, and independent of the width of the aperture. The constancy of the approach speed suggests a simple means by which optic flow can be used to gauge the distance and width of the aperture, and guide wing closure.
Flying animals accomplish high-speed navigation through fields of obstacles using a suite of sensory modalities that blend spatial memory with input from vision, tactile sensing, and, in the case of most bats and some other animals, echolocation. Although a good deal of previous research has been focused on the role of individual modes of sensing in animal locomotion, our understanding of sensory integration and the interplay among modalities is still meager. To understand how bats integrate sensory input from echolocation, vision, and spatial memory, we conducted an experiment in which bats flying in their natural habitat were challenged over the course of several evening emergences with a novel obstacle placed in their flight path. Our analysis of reconstructed flight data suggests that vision, echolocation, and spatial memory together with the possible exercise of an ability in using predictive navigation are mutually reinforcing aspects of a composite perceptual system that guides flight. Together with the recent development in robotics, our paper points to the possible interpretation that while each stream of sensory information plays an important role in bat navigation, it is the emergent effects of combining modalities that enable bats to fly through complex spaces.
- Philosophical transactions of the Royal Society of London. Series B, Biological sciences
- Published over 2 years ago
Insect- and bird-size drones-micro air vehicles (MAV) that can perform autonomous flight in natural and man-made environments are now an active and well-integrated research area. MAVs normally operate at a low speed in a Reynolds number regime of 10(4)-10(5) or lower, in which most flying animals of insects, birds and bats fly, and encounter unconventional challenges in generating sufficient aerodynamic forces to stay airborne and in controlling flight autonomy to achieve complex manoeuvres. Flying insects that power and control flight by flapping wings are capable of sophisticated aerodynamic force production and precise, agile manoeuvring, through an integrated system consisting of wings to generate aerodynamic force, muscles to move the wings and a control system to modulate power output from the muscles. In this article, we give a selective review on the state of the art of biomechanics in bioinspired flight systems in terms of flapping and flexible wing aerodynamics, flight dynamics and stability, passive and active mechanisms in stabilization and control, as well as flapping flight in unsteady environments. We further highlight recent advances in biomimetics of flapping-wing MAVs with a specific focus on insect-inspired wing design and fabrication, as well as sensing systems.This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’.
Dragonfly has excellent flight performance and maneuverability due to the complex vein structure of wing. In this research, nodus as an important structural element of the dragonfly wing is investigated through an experimental visualization approach. Three vein structures were fabricated as, open-nodus structure, closed-nodus structure (with a flex-limiter) and rigid wing. The samples were conducted in a wind tunnel with a high speed camera to visualize the deformation of wing structure in order to study the function of nodus structured wing in gliding flight. According to the experimental results, nodus has a great influence on the flexibility of the wing structure. Moreover, the closed-nodus wing (with a flex-limiter) enables the vein structure to be flexible without losing the strength and rigidity of the joint. These findings enhance the knowledge of insect-inspired nodus structured wing and facilitate the application of Micro Air Vehicle (MAV) in gliding flight.