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Concept: Wing


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.

Concepts: Bird flight, Geometry, Flight, Jurassic, Archaeopteryx, Wing, Dinosaur, Bird


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.

Concepts: Force, Mass, Insect, Flying and gliding animals, Bat, Flight, Wing, Classical mechanics


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.

Concepts: Lizard, Aerodynamics, Airfoil, Leading edge slats, Flight, Fixed-wing aircraft, Flying and gliding animals, Wing


Competition is one of the most cited mechanisms to explain secondary sexual dimorphism in animals. Nonetheless, it has been proposed that sexual dimorphism in bat wings is also a result of adaptive pressures to compensate additional weight caused by fetus or pup carrying during the reproductive period of females. The main objective of this study is to verify the existence of sexual dimorphism in Sturnira lilium wings. We employed geometric morphometrics techniques using anatomical landmarks superimposition to obtain size (Centroid Size) and shape variables of wings, which were reduced by Linear Discriminant Analysis (LDA). We also employed classical morphometrics using wing length measurements to compare efficiency between these two morphometric approaches and make comparisons using wing area measurements. LDA indicated significant differences between wing shapes of males and females, with 91% (stepwise classification) and 80% (leave-one-out cross validation) of correct classification. However, the size variable obtained did not contribute to such classifications. We have observed larger areas in female wings, but we found no differences in wing length measurements and no allometric effects in wing length, shape and area measurements. Interestingly, our study has provided evidences of morphological differences where classical morphometrics have failed. LDA and area measurements analyses revealed that females have a different area distribution in distinct portions of the wing, with wider dactylopatagia and plagiopatagia, and wingtips more triangular than males. No differences in body length or relative wing length were observed between the sexes, but pregnant females have more body weight than non-pregnant females and males. Our findings suggest that sexual dimorphism in the wing shape of S. lilium is probably related to the increase in flight efficiency of females during reproductive period. It decreases wing loading in specific portions of the wing and reduces energy cost to maintain a faster and maneuverable flight.

Concepts: Reproduction, Morphometrics, Wing, Female, Gender, Sexual dimorphism, Sex, Male


BACKGROUND: Anopheles (Kerteszia) cruzii is a primary vector of Plasmodium parasites in Brazil’s Atlantic Forest. Adult females of An. cruzii and An. homunculus, which is a secondary malaria vector, are morphologically similar and difficult to distinguish when using external morphological characteristics only. These two species may occur syntopically with An. bellator, which is also a potential vector of Plasmodium species and is morphologically similar to An. cruzii and An. homunculus. Identification of these species based on female specimens is often jeopardised by polymorphisms, overlapping morphological characteristics and damage caused to specimens during collection. Wing geometric morphometrics has been used to distinguish several insect species; however, this economical and powerful tool has not been applied to Kerteszia species. Our objective was to assess wing geometry to distinguish An. cruzii, An. homunculus and An. bellator. METHODS: Specimens were collected in an area in the Serra do Mar hotspot biodiversity corridor of the Atlantic Forest biome (Cananeia municipality, State of Sao Paulo, Brazil). The right wings of females of An. cruzii (n= 40), An. homunculus (n= 50) and An. bellator (n= 27) were photographed. For each individual, 18 wing landmarks were subjected to standard geometric morphometrics. Discriminant analysis of Procrustean coordinates was performed to quantify wing shape variation. RESULTS: Individuals clustered into three distinct groups according to species with a slight overlap between representatives of An. cruzii and An. homunculus. The Mahalanobis distance between An. cruzii and An. homunculus was consistently lower (3.50) than that between An. cruzii and An. bellator (4.58) or An. homunculus and An. bellator (4.32). Pairwise cross-validated reclassification showed that geometric morphometrics is an effective analytical method to distinguish between An. bellator, An. cruzii and An. homunculus with a reliability rate varying between 78-88%. Shape analysis revealed that the wings of An. homunculus are narrower than those of An. cruzii and that An. bellator is different from both of the congeneric species. CONCLUSION: It is possible to distinguish among the vectors An. cruzii, An. homunculus and An. bellator based on female wing characteristics.

Concepts: Wing, Biodiversity, Procrustes analysis, São Paulo, Malaria, Plasmodium, Plasmodium falciparum, Brazil


BACKGROUND: Wing size and shape have important aerodynamic implications on flight performance. We explored how wing size was related to wing shape in territorial males of 37 taxa of the damselfly family Calopterygidae. Wing coloration was also included in the analyses because it is sexually and naturally selected and has been shown to be related to wing shape. We studied wing shape using both the non-dimensional radius of the second moment of wing area (RSM) and geometric morphometrics. Lower values of the RSM result in less energetically demanding flight and wider ranges of flight speed. We also re-analyzed previously published data on other damselflies and dragonflies. RESULTS: The RSM showed a hump-shaped relationship with wing size. However, after correcting for phylogeny using independent contrast, this pattern changed to a negative linear relationship. The basal genus of the study family, Hetaerina, was mainly driving that change. The obtained patterns were specific for the study family and differed from other damselflies and dragonflies. The relationship between the RSM and wing shape measured by geometric morphometrics was linear, but relatively small changes along the RSM axis can result in large changes in wing shape. Our results also showed that wing coloration may have some effect on RSM. CONCLUSIONS: We found that RSM showed a complex relationship with size in calopterygid damselflies, probably as a result of other selection pressures besides wing size per se. Wing coloration and specific behavior (e.g. courtship) are potential candidates for explaining the complexity. Univariate measures of wing shape such as RSM are more intuitive but lack the high resolution of other multivariate techniques such as geometric morphometrics. We suggest that the relationship between wing shape and size are taxa-specific and differ among closely-related insect groups.

Concepts: Moment, Damselflies, Dragonfly, Change, Wing, Odonata, Aerodynamics, Damselfly


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.

Concepts: Optics, Aperture, Flight, Flying and gliding animals, Denny Laine, Bird flight, Bird, Wing


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.

Concepts: Understanding, Wing, Sense, Flight, Sensory system, Insect, Bat, Flying and gliding animals


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’.

Concepts: Aerospace engineering, Wing, Bat, Drag, Flight, Insect, Flying and gliding animals, Aerodynamics


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.

Concepts: Bird flight, System, Wing, Aerodynamics, Aircraft, Flying and gliding animals, Structure, Flight